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BERGEY'S MANUAL 

OF 

DETERMINATIVE 
BACTERIOLOGY 



BY 

ROBERT S. BREED 

New York State Experiment Station {Cornell University), Geneva, New York 

E. G. D. MURRAY^ 

McGill University, Montreal, Province Quebec, Canada 

A. PARKER HITCHENS 

University of Pennsylvania, Philadelphia, Pennsylvania 

Assisted by 

Sixty Contributors Whose Names and Contributions Appear 

in the Pages Immediately Following 



SIXTH EDITION 



fi/i{irsA^s\\si\ 

iKVAVTACHE j'At 



BALTEVIORE 

THE WILLIAMS & WILKINS COMPANY 
1948 



First Edition, August, 1923 

Second Edition, December, 1925 

Third Edition, January, 1930 

Fourth Edition, March, 1934 

Preprint of pages ix + 79 of Fifth Edition, October, 1938 

Fifth Edition, April, 1939 



Copyright, 1948 
The Williams & Wilkins Company 

Made in United States of America 
Published January, 1948 



Composed and Printed at thb 
WAVERLY PRESS, INC. 

FOS 

The Williams & Wilkins Compajjy 

Mt. Royal and Guilford Aves. 

Baltimore, Md., U. S. A. 



LIST OF CONTRIBUTORS 



Allen, O. N. 
Baldwin, I. L. 
Barker, H. A. 



Bengtson, Ida A. 
Bergey, D. H.f 



Borman, Earl L. 
Branham, Sara E. 
Breed, Robert S. 



Robert S. and Margaret 

E. Breed 
Buchanan, R. E. 



Burkholder, Walter H. 



Rhizobium 223 

Rhizobium 223 

Methanococcus 248 

Butyribacterium 380 

Methanobacterium 645 

Rickeitsiaceae 1083 

Bartonellaceae 1100 

Methanomonas 179 

Mycoplana 191 

Thiospira 212 

Achromobacter 417 

Flavobacterium 427 

Dialister 594 

Paracolobactrum 460 

N eisseriaceae 295 

Survey of Classifications 5-38 

Rules of Nomenclature 49-64 

Nitrobacteriaceae 69 

Pseudomonadaceae 62, 171 

Chromobacteriurn 231 

Methanococcus 248 

Pediococcus 249 

Sarcina 285 

Leptotrichia 264 

Corynebacteriaceae 381 

Achromobacteriaceae 412 

App. 2. Eschericheae 461 

Serratia 479 

Malleomyces 554 

Fusobacterium, 581 and Fusiformis 583 

Bacteriaceae 596 

Caulobacteriineae 827 

Chlamydobacteriales 981 

Indexes 1297 

How Bacteria Are Named and Identified 39-48 

Etymology 64 ff. 

Myxobacteriales 1005 

Pseudomonadaceae 82 and 150 

Corynebacterium 381 

Erwima 463 

Bacterium 638 



t Deceased, September, 1937. 



IV 



LIST OF CONTRIBUTORS 



Chapman, Orren D. Klebsiella 457 

Donovania 559 

Chester, F. D.f Erwinia 463 

Bacillus 704 

Clise, Eleanore H. App. 3. Micrococcus 252 

Pasteurella 546 

App. 1. Bacteroides, 575; App. Eubacteriineae, 692; 
App. Nocardia, 915; App. Streptomyces, 967; and 
App. Spirochaetales, 1051 

Conn, H. J. Nitrobacteriaceae 69 

Agrobacterium 227 

App. 3. Corynebacterium 407 

Alcaligenes 412 

Davis, Gordon E. Borrelia 1058 

Dienes, Louis Borrelomyceiaceae 1287 

Edwards, P. R. Salmonella 492 

Evans, Alice C. Parvobacteriaceae 545 

Fred, E. B. Rhizobium 223 

Hagan, W. A. Parvobacteriaceae 545 

Hall, Ivan C. Anaerobic section Micrococcus, 246; Neisseria, 255; 

Veillonella, 302; Diplococcus, 308; and Strepto- 
coccus, 328 

Hanks, John H. Mycobacterium 876 

Harvey, Philip Pasteurella 546 

Haynes, Wm. C. Vibrio 192 

Spirillum 216 

Henrici, A. T.J Caulobacieriineae 827 

Nocardia 892 

Actinomyces 925 

Streptomyces 929 

Chlamydobacteriales 981 

Hitchens, A. Parker Vibrio 192 

Mimeae 595 

Editor Supplement I, 1082, II, 1125 and III, 1287 

Hitchner, E. R. Aeromonas 101 

Hofer, A. W. Azotobacleriaceae 219 

Holmes, Francis O. Virales 1200 

Hucker, G.J. Micrococcus 235 

Gaffkya 283 

Streptococcus 312 

Leuconostoc 346 

Huddleson, I. F. Brucella 560 

Johnson, Frank H. Phosphorescent Bacteria 633 

Kauffmann, F. Salmonella 492 

Kelly, C. D. Acetobacler 179 

Bacteroides 564 

Noguchia 592 

Kirby, Harold App. 1. Rickettsiales 1121 

Meyer, K. F. Pasteurella 545 

t Deceased, January, 1943. J Deceased, April, 1943. 



LIST OF CO>n'RIBUTORS 



Murray, E. G. D. 



Pederson, C. S. 



Peshkoff, Michael A. 
Pittman, Margaret 
Rahn, Otto 
Rake, Geoffrey 
Reed, Guilford B. 
Rettger, L. F. 
Robinson, George H.t 
Roy, T. E. 
Rustigian, Robert 
Sherman, J. M. 

Smith, Frederick 

Smith, X. R. 
Speck, M. L. 
Spray, R. S. 
Stanier, R. Y. 

Starkey, R. L. 
Steinhaus, Edward A. 

Stuart, C. A. 
Tobie, W. C. 
Van Xiel, C. B. 



Vaughn, Reese 
Waksman, Selman A. 

Weiimian, David 
Welsh, Mark 
Yale, M. W. 



ZoBell, Claude E. 



Neisseriaceae 295 

Diplococcus 305 

Streptococcus 312 

Corynebacteriaceae 381 

Parvobacteriaceae 545 

Spirochaetales 1051 

Leuconostoc 346 

Lactobacillus 349 

Lepiotrichia 364 

Microbacterium 370 

Butyribacterium 380 

Caryophanales 1002 

Hemophilus 584 

Characterizations of Groups 64 ff. 

Chlamydozoaceae 1114 

Mycobacteriaceae 875 

Lactobacillus 349 

Spirochaetales 1051 

Bacteroides 564 

Proteus 486 

Streptococcus 312 

Lactobacillus 349 

Salmonella 492 

Sh igella 525 

Bacillus 704 

Microbacterium 370 

Clostridium 763 

Cytophagaceae 1012 

Sporocytophagaceae 1048 

Xitrobacteriaceae 69 

Insect Microbiology 417, 742 ff. 

Rickettsiaceae 1083 

Proteus 486 

Chromobacterium 231 

Propionibacterium 372 

Rhodobacteriineae 838 

Bcggiatoaceae 988 

Achromatiaceae 997 

Acetobacter 179 

Actinomycetaceae 892 

Streptomycetaceae 929 

Bartonellaceae 1100 

Pasteurella 546 

Escherichia 444 

Aerobacter 453 

Proteus 486 

Desulfovibrio 209 

Marine Microbiology 107, 418, 431 ff. 



t Deceased, October, 1945. 



PREFACE TO SIXTH EDITION 

More than the usual amount of time and effort has been given toward 
making this new edition of Bergey's Manual useful. The volume has 
been completely revised and is reset in double column format so that each 
page carries about 20 per cent more type than the pages in the fifth edition. 
Those who are interested in special groups of bacteria will find something 
new in the presentation of the relationships in every genus. Because 
of our rapidly expanding knowledge, changes in the outline classification 
and text were made necessary. These changes have in every case been 
made by speciahsts in consultation with the Editorial Board. Every spe- 
cialist possesses first hand knowledge of the species in the group that he or 
she has reviewed. 

Because increasing knowledge has shown the fission fungi to be a larger 
and more diversified group than previously realized, the number of species 
described has increased from 1335 in the fifth edition to 1630 in the present 
edition of the Manual. This number does not cover all of the descriptions 
found in the literature for, as in all other fields of biology, many of the 
descriptions are so inadequate that the species described cannot now be 
identified. Many descriptions are obvious or probable duplications of 
previous descriptions while still others are based on nothing more substan- 
tial than the author's belief that he had something new, he having made 
but little effort to compare his cultures with those found by previous 
investigators. An indication of the large number of inadequate descrip- 
tions will be found by referring to the material in the appendixes to the 
various groups, and to the index where synonyms and incompletely 
described species are shown in italics. 

The large number of these poorly described species suggests that there 
has been much unsatisfactory work done in the field of bacteriological 
taxonomy. Progress in this inadequately developed field is needed as it 
would help to clarify the approach to desirable research in many fields of 
bacteriology. 

It is believed that both teachers and investigators will find the new 
Source and Habitat index useful. It is important to know what organisms 
have been described from any given habitat in determining the identity of 
a described species or whether a given species is new. 

The future development of taxonomic work holds several interesting 
possibilities of increased international cooperation such as between the 
various National Type Culture Collections and within the International 



Vm PKEFACE TO SIXTH EDITION 

Association of Microbiologists. The Trust Funds provided through the 
generosity of Dr. Bergey before his death have been used in developing the 
present edition of the Manual and future funds are to be used in the same 
way under the management of a self-perpetuating Board of Editor-Trustees. 

We are all under obligation to those who have given so freely of their 
time and special knowledge in preparing this edition cf the Manual. 
Moreover the Editor-in-Chief is under special obligation to his wife, 
Margaret Edson Breed who has carried the burden of the indexing; to Mrs. 
Eleanore Heist Clise who has given invaluable service in bibliographical 
research, in proof reading and other ways; and to his secretary, Miss Maude 
Hogan, who has cared for many difficult manuscripts and a voluminous 
technical correspondence. 

Many binomials not previously mentioned in the Manual will be found 
in the Index of Genus and Species Names. Each new name means that 
there is a new bibliographic reference in the text. Practically all of the 
incomplete references of previous editions and all new references have been 
examined in the original, something that is essential in all accurate taxo- 
nomic work. The index of names is the most complete list that has ap- 
peared in the literature and should always be consulted before new genus 
or species names are proposed. 

This edition of the Manual has been more than four years in press, thanks 
to the care that has been taken to make it complete and useful. Through- 
out, the Editorial Board has had the cooperation and understanding help 
of the publishers of the book who themselves have been forced to meet and 
overcome the trying difficulties of the war years. 

The plan of the present book is such that it will be found useful both to 
teachers and research workers. 

Robert S. Breed, Chairman 
E. G. D. Murray 
A. Parker Kitchens 
Board of Editor-Trustee.?. 
April, 1947. 



PREFACE OF FIRST EDITION 

The elaborate system of classification of the bacteria into families, tribes 
and genera by a Committee on Characterization and Classification of the 
Society of American Bacteriologists (1917, 1920) has made it very desirable 
to be able to place in the hands of students a more detailed key for the 
identification of species than any that is available at present. The valuable 
book on "Determinative Bacteriologj'^" by Professor F. D. Chester, pub- 
lished in 1 901 , is now of very little assistance to the student, and all previous 
classifications are of still less value, especially as earlier systems of classifica- 
tion were based entirely on morphologic characters. 

It is hoped that this manual will serve to stimulate efforts to perfect the 
classification of bacteria, especially by emphasizing the valuable features 
as well as the weaker points in the new system which the Committee of the 
Society of American Bacteriologists has promulgated. The Committee 
does not regard the classification of species offered here as in any sense 
final, but merel}^ a progress report leading to more satisfactory classification 
in the future. 

The Committee desires to express its appreciation and thanks to those 
members of the society who gave valuable aid in the compilation of material 
and the classification of certain species. . . . 

The assistance of all bacteriologists is earnestly solicited in the cori-ection 
of possible errors in the text; in the collection of descriptions of all bacteria 
that may have been omitted from the text; in supplying more detailed 
descriptions of such organisms as are described incompletely ; and in furnish- 
ing complete descriptions of new organisms that maj' be discovered, or 
in directing the attention of the Committee to publications of such newly 
described bacteria. 

David H. Bergey, Chairman 
Francis C. Harrison 
Robert S. Breed 
Bernard W. Hammer 
Frank M. Huntoon 
Committee on Manual. 
August, 1923. 



.'^AS^J 



■H 



CONTENTS 

Introduction 1 

Historical Survey of Classifications 5 

How Bacteria are Named and Identified 39 

Rules of Nomenclature 49 

Class Schizomyceies Nageli 65 

Order I. Eubacteriales Buchanan 66 

Suborder I. Eubacteriineae Breed, Murray and Hitchens 67 

Family I. Nitrobacteriaceae Buchanan 69 

Tribe I. Niirobacterieae Winslow et al 70 

Genus I. Nitrosomoyias Winogradsky 70 

Genus II. Nilrosococcus Winogradsky 71 

Genus III. Nitrosospira Winogradsky 71 

Genus IV. Nitrosocystis Winogradsky 72 

Genus V. Nitrosogloea H. Winogradsky 73 

Genus VI. Nitrobacter Winogradsky 74 

Genus VII. Nitrocystis H. Winogradsky 75 

Tribe II. Hydrogenomonadeae Pribram 76 

Genus I. Hydrogenomonas Orla-Jensen 76 

Tribe III. Thiobacilleae Bergey, Breed and Murray 78 

Genus I. Thiobacillus Beijerinck 78 

Family II. Pseudomonadaceae Winslow et al 82 

Tribe I. Pseudomonadeae Kluyver and Van Niel 82 

Genus I. Pseudomonas Migula 82 

Genus II. Xanthomonas Dowson 150 

Genus III. Methanomonas Orla-Jensen 179 

Genus IV. Acetobacter Beijerinck 179 

Genus V. Protaminobacter den Dooren de Jong 189 

Genus VI. Mycoplana Gray and Thornton 192 

Tribe II. Spirilleae Kluyver and Van Niel 192 

Genus I. Vibrio Miiller 192 

Genus II. Desulfovibrio Kluyver and Van Niel 207 

Genus III. Cellvibrio Winogradsky 209 

Genus IV. Cellfalcicula Winogradsky 211 

Genus V. Thiospira Vislouch 212 

Genus VI. Spirillum Ehrenberg 212 

Family III. Azotobacteriaceae Bergey, Breed and Murray 219 

Genus I. Azotobacter Beijerinck 219 

Appendix : Genus A. Azotomonas Stapp 221 

Family IV. Rhizobiaceae Conn 223 

Genus I. Rhizobium Frank 223 

Genus II. Agrobacterium Conn 227 

, Genus III. Chromobacterium Bergonzini 231 

Family V. Micrococcaceae Pribram 235 

Genus I. Micrococcus Cohn 235 

Appendix: Genus A. Methanococcus Kluyver and Van Niel 248 

Genus B. Pediococctis Balcke 249 



XI 



7ij 



Xll CONTENTS 

Genus II. Gaffkya Trevisan 283 

Genus III. Sarcina Goodsir 285 

Subgenera: 

Zymosarcina Smit 285 

Methanosarcina Kluy ver and Van Niel 285 

Sarcinococcus Breed 285 

Sporosarcina Orla-Jensen 285 

Family VI. Neisseriaceae Pr^vot 295 

Genus I. Neisseria Trevisan 295 

Genus II. Veillonella Prevot 302 

Family VII. Lactobacteriaceae Orla-Jensen 305 

Tribe I. Slreptococceae Trevisan 305 

Genus I. Diplococcus Weichselbaum 305 

Genus II. Streptococcus Rosenbach 312 

Genus III. Leuconostoc Van Tieghem 346 

Tribe II. Lactobacilleae Winslow et al 349 

Genus I. Lactobacillus Beijerinck 349 

Sub-genera: 

Thermobactcrivii) Orla-Jensen 350 

Streptobacterium Orla-Jensen 350 

Betabacterium Orla-Jensen 350 

Appendix: Genus A. Leptotrichia Trevisan 365 

Genus II. Microbacierium Orla-Jensen 370 

Genus III. Propionibacterium Orla-Jensen 372 

Genus IV. Butyribacterium Barker 379 

Family VIII. Corynebacteriaceae Lehmann and Neumann 381 

Genus I. Corynebacterium Lehmann and Neumann 381 

Genus II. Listeria Pirie 408 

Genus III. Erysipelothrix Rosenbach 410 

Family IX. Achromobacteriaceae 412 

Genus I. Alkaligenes Castellani and Chalmers 412 

Genus II. Achromobacter Bergey et al 417 

Genus III. Flavobacterium Bergey et al 427 

Family X. Enterobacteriaceae Rahn 443 

Tribe I. Eschericheae Bergey, Breed and Murray 444 

Genus I. Escherichia Castellani and Chalmers 444 

Genus II. Aerobacter Beijerinck 453 

Genus III. Klebsiella Trevisan 457 

Appendix: Genus A. Paracolobactruni Borman, Stuart and 

Wheeler 459 

Tribe II. Erwineae Winslow et al 463 

Genus I. Erwinia Winslow et al 463 

Tribe III. Serrateae Bergey, Breed and Murray 479 

Genus I. Serratia Bizio 3mend. Breed and Breed 479 

Tribe IV. Proteae Castellani and Chalmers 486 

Genus I. Proteus Hauser 486 

Tribe V. Salmonelleae Bergey, Breed and Murray 492 

Genus I. Salmonella Lignieres 492 

Genus II. Shigella Castellani and Chalmers 535 



CONTENTS XIU 

Family XI. Parvobacteriaceae Rahn 545 

Tribe I. Pasteurelleae Castellani and Chalmers 545 

Genus I. Pasteurella Trevisan 546 

Genus II. Malleomyces Hallier 554 

Genus III. Actinobacillus Brumpt 556 

Appendix: Genus A. Donovania Anderson et al 558 

Tribe II. Brucellcae Bergey, Breed and Murray 560 

Genus I. Brucella Meyer and Shaw 560 

Tribe III. Bacteroideae Breed, Murray and Kitchens 564 

Genus I. Bacteroides Castellani and Chalmers 564 

Genus II. Fusohacterium Knorr 581 

Appendix : Genus A. Fusiformis Hoelling 583 

Tribe IV. Hemophileae Winslow et al 584 

Genus I. Hemophilus Winslow et al 584 

Genus II. Moraxella Lwoff 590 

Genus III. Noguchia Olitsky, Sy vert on and Tyler 592 

Genus IV. Dialister Bergey et al 594 

Appendix : Tribe Mimeae DeBord 595 

Family XII. Bacteriaceae Cohn 596 

Genus I. Bacterium Ehrenberg 596 

Subgenera: 

Kurlhia Trevisan 600 

Cellulomonas Bergey et al 613 

Saccharobacterium Sickles and Shaw 623 

Agarbacterium Angst 624 

Photobacterium Beijerinck 633 

Methanobacterium Kluyver and Van Niel 645 

Appendix: Suborder Eubacteriineae: Overlooked species and syn- 
onyms 692 

Family XIII. Bacillaceae Fisher 704 

Genus I. Bacillus Cohn 705 

Genus II. Clostridium Prazmowski 763 

Suborder II. Caulobacteriineae Breed, Murray and Kitchens 828 

Family I. Nerskiaceae Kenrici and Johnson 830 

Genus I. Nevskia Famintzin 830 

Family II. Gallionellaceae Kenrici and Johnson 830 

Genus I. Gallionella Ehrenberg 831 

Family III. Caulobacteriaceae Kenrici and Johnson 832 

Genus I. Caulobacter Kenrici and Johnson 832 

Family IV. Siderocapsaceae Pribram , . 833 

Genus I. Siderocapsa Molisch 833 

Genus II. Sideromonas Cholodny 834 

Appendix: Family Pasteuriaceae Laurent 836 

Genus I. Pasteuria Metchnikoff 836 

Genus II. Blastocaulis Kenrici and Johnson 836 

Suborder III. Rhodobacteriineae Breed, Murray and Kitchens 838 

Family I. Thiorhodaceae Molisch 841 

Genus I. Thiosarcina Winogradsky 842 

Genus II. Thiopedia Winogradsky 843 

Genus III. Thiocapsa Winogradsky 844 

Genus IV. Thiodictyon Winogradsky 845 



XIV CONTENTS 

Genus V. Thiothece Winogradsky 846 

Genus VI. Thiocystis Winogradsky 846 

Genus VII. Lamprocystis Schroeter 847 

Genus VIII. Amoebobacter Winogradsky 848 

Genus IX. Thiopolycoccus Winogradsky 860 

Genus X. Thiospirillum Winogradsky 850 

Genus XI. Rhabdomonas Cohn 853 

Genus XII. Rhodothece Molisch 855 

Genus XIII. Chromatium Perty 856 

Family II. Aihiorhodaceae Molisch 861 

Genus I. Rhodopseudomonas Kluyver and Van Niel emend. Van 

Niel 861 

Genus II. Rhodospirillum Molisch 866 

Family III. Chlorobacteriaceae Geitler and Pascher 869 

Genus I. Chlorobium Nadson 869 

Genus II. Pelodictyon Lauterborn 870 

Genus III. Clathrochloris Geitler 872 

Genus IV. Chlorobacterium Lauterborn 872 

Genus V. Chlorochromatium Lauterborn 873 

Genus VI. Cylindrogloea Perfiliew 873 

Order II. Actinomycetales Buchanan 875 

Family I. Mycobacteriaceae Chester 875 

Genus I. Mycobacterium Lehmann and Neumann 876 

Family II. Actinomyceiaceae Buchanan 892 

Genus I. Nocardia Trevisan 892 

Genus II. Actinomyces Harz 925 

Family III. Streptomycetaceae Waksman and Henrici 929 

Genus I. Streptomyces Waksman and Henrici 929 

Genus II. Micromonospora Prskov 978 

Order III. Chlamydobacteriales Buchanan 981 

Family I. Chlamydobacteriaceae Migula 981 

Genus I. Sphaerotilus Kiitzing 982 

Genus II. Clonothrix Roze 983 

Genus III. Leptothrix Ktitzing 983 

Family II. Crenothricaceae Hansgirg 987 

Genus I. Crenothrix Cohn 987 

Family III. Beggiatoaceae Migula 988 

Genus I. Thiothrix Winogradsky 988 

Genus II. Beggiatoa Trevisan 990 

Genus III. Thiospirillopsis Uphof 993 

Genus IV. Thioploca Lauterborn 993 

Appendix: Family Achromatiaceae Massart 997 

Genus I. Achromaiium Schewiakoff 997 

Genus II. Thiovulum Hinze 999 

Genus III. Macromonas Utermohl and Koppe 1000 

Appendix : Order Caryophanales Peshkoff 1002 

Family I. Pontothricaceae Peshkoff 1002 

Genus I. Pontothrix Nadson and Krassilnikow 1002 

Family II. Arthromitaceae Peshkoff 1002 

Genus I. Arthromitus Leidy 1002 

Genus II. Coleomitus Duboscq and Grass^ 1003 



CONTENTS XV 

Family III. Oscillospiraceae Peshkoff 1003 

Genus I. Oscillospira Chatton and Perard 1004 

Family IV. Caryophanaceae Peshkoff 1004 

Genus I. Caryophanon Peshkoff 1004 

Order IV. Myxobacteriales Jahn 1005 

Family I. Cytophagaceae Stanier 1012 

Genus I. Cytophaga Stanier 1012 

Family II. Archangiaceae Jahn 1017 

Genus I. Archangium Jahn 1017 

Genus II. Stelangium Jahn 1020 

Family III. Sorangiaceae Jahn 1021 

Genus I. Sorangium Jahn 1021 

Familj' IV. Polyangiaceae Jahn 1025 

Genus I. Polyangium Jahn 1025 

Genus II. Synangium Jahn 1032 

Genus III. Melittangium Jahn 1033 

Genus IV. Podangium Jahn 1034 

Genus V. Chondromyces Berkeley and Curtis 1036 

Family V. Mycococcaceae Jahn 1040 

Genus I. M yxococcus Thaxter 1040 

Genus II. Chrondrococcus Jahn 1044 

Genus III. Angiococcus Jahn 1047 

Genus IV. Sporocyfophaga Stanier 1048 

Order V. Spirochaetales Buchanan 1051 

Family I. Spirochaetaceae Swellengrebel 1051 

Genus I. Spirochaeta Ehrenberg 1051 

Genus II. Saprospira Gross 1054 

Genus III. Cristispira Gross 1055 

Family II. Treponemataceae Schaudinn 1058 

Genus I. Borrelia Swellengrebel 1058 

Genus II. Treponema Swellengrebel 1071 

Genus III. Leptospira Noguchi 1076 

Supplement 1 1081 

Order Rickettsiales Gieszcykiewicz 1083 

Family I. Rickettsiaceae Pinkerton 1083 

Genus I. Rickettsia da Rocha-Lima 1084 

Genus II. Coxiella Bengtson 1092 

Genus III. Cowdria Bengtson 1094 

Family II. Bartonellaceae Gieszcykiewicz 1100 

Genus I. Bartonella Strong, Tyzzer and Sellards 1100 

Genus II. Haemohartonella Tyzzer and Weinman 1102 

Genus III. Grahamella Brumpt 1109 

Genus IV. Eperythrozoon Schilling 1111 

Family III. Chlamydozoaceae Moshkovskj^ 1114 

Genus I. Chlamydozodn Halberstaedter and von Prowazek 1114 

Genus II. Miyagawanella Brumpt 1115 

Genus III. Colesiota Rake 1119 

Appendix : Genus A. Caryococcus Dangeard 1121 

Genus B. Drepanospira Petschenko 1122 

Genus C. Holospora Haffkine 1122 



XVI CONTENTS 

Supplement II 1125 

Order Virales Breed, Murray and Kitchens , 1128 

Suborder 1. Phagineae Holmes 1128 

Family I. Phagaceae Holmes 1128 

Genus I. Phagus Holmes 1128 

Suborder II. Phytophagineae Holmes 1145 

Family I. Chlorogenaceae Holmes 1145 

Genus I. Chlorogenus Holmes 1146 

Genus II. Carpophthora McKinney 1151 

Genus III. Morsus Holmes 1153 

Genus IV. Aureogenus Black 1154 

Genus V. Galla Holmes 1157 

Genus VI. Fractilinae McKinney 1159 

Family II. Marmoraceae Holmes 1163 

Genus I. Marmor Holmes 1 163 

Genus II. Acrogenus Holmes 1202 

Genus III. Corium Holmes 1203 

Genus IV. Nanus Holmes 1206 

Genus V. Rimocortius Milbrath and Zeller 1208 

Genus VI. Adelonosus Brierley and Smith 1211 

Family III. Annulaceae Holmes 1212 

Genus I. Annulus Holmes 1212 

Family IV. Rugaceae Holmes 1218 

Genus I. Ruga Holmes 1218 

Family V. Savoiaceae Holmes 1221 

Genus I. Savoia Holmes 1221 

Family VI. Lethaceae Holmes 1223 

Genus I. Lethum Holmes 1223 

Suborder III. Zoophagineae Holmes 1225 

Family I. Borrelinaceae Holmes 1225 

Genus I. Borrelina Holmes 1225 

Genus II. Moralor Holmes 1227 

Family II. Borreliotaceae Holmes 1229 

Genus I. Borreliota Goodpasture 1229 

Genus II. Briareus Holmes 1233 

Genus III. Scelus Holmes 1234 

Genus IV. Hostis Holmes 1239 

Genus V. Moliler Holmes 1241 

Family III. Erronaceae Holmes 1248 

Genus I. Erro Holmes 1248 

Genus II. Legio Holmes 1257 

Genus III. Formido Holmes 1263 

Family IV. Charonaceae Holmes 1265 

Genus I. Charon Holmes 1265 

Genus II. Tarpeia Holmes 1268 

Genus III. Tortor Holmes 1275 

Family V. Trifuriaceae Holmes 1282 

Genus I. Trifur Holmes 1282 

Family VI. Rabulaceae Holmes 1284 

Genus I. Rabula Holmes 1284 

Supplement III 1287 

Family Borrelomycetaceae Turner 1291 

Genus I. Ascococcus Borrel et al 1291 



INTRODUCTION 

Suggestions for the Use of the "Manual in Classifying 
Unknown Organisms 

No organism can be classified before we have determined, through de- 
tailed study, its morphological, cultural, physiological and pathogenic 
characters. 

The characters used in the keys to orders, families and genera may ordi- 
narily be determined by the use of a dozen or more of the procedures de- 
scribed in the ^vlanual of Pure Culture Study issued by the Committee on 
Bacteriological Technic (H. J. Conn, Chairman, Geneva, New York) of the 
Society of American Bacteriologists. j\Iore complete examinations must 
be made as indicated in the iNIanual of Pure Culture Study, and in the 
Descriptive Charts which accompany this Manual where it is desired to 
identify individual species. These tests must be made if bacteria are to be 
accurately identified and described. 

It is urged that beginning students be taught the technics necessary for 
the identification of species in the hope that the taxonomic work of the 
future may be placed on a more satisfactory basis. 

After a complete study of the characters of the organism has been 
made, turn to page 65 and ascertain first in which order the organism 
belongs. When the order and suborder (if necessary) have been ascer- 
tained, turn to the page of the IManual on which the key to that order or 
suborder is given. In this key ascertain the family or subfamily to which 
the organism belongs. 

When the family or subfamily has been decided on, again refer to the 
page of the Manual on which the key to that familj^ or subfamily is given. 
In this key ascertain the tribe to which the organism belongs. 

When the tribe has been decided on, again find the page of the Manual 
on which the key to the tribe is given. In this key ascertain the genus 
to which the organism belongs. 

When the genus has been decided on, again refer to the page of the 
Manual on which the key to that genus is given. In this key, trace out 
the species under investigation. 

For example, if one wishes to trace a short, peritrichous, Gram-negative, 
non-spore-forming rod that grows well on ordinary culture media at 37°C, 
fermenting glucose and lactose with production of acid and gas, not 
liquefying gelatin, producing no pigment on any culture medium, with 
negative reaction for acetylmethylcarbinol, producing indole and reducing 
nitrates, consult the key to the orders on page 65. 

1 



2 SUGGESTIONS FOR USE OF MANUAL 

In this key examine A. Cells rigid, not flexuous. This indicates our or- 
ganism as its cells remain constant in form. 

We next examine 1. Cells single, in chains or masses. Not branching and 
mycelial in character. Not arranged in filaments. Not acid fast. As the 
organism in question occurs as single cells or at most as short chains and is 
not acid fast, this indicates that it belongs to the Order Eubacteriales. 

We now examine a. Do not possess -photo synthetic pigments. Cells do not 
contain free sulfur. As our organism is unpigmented and the cells do not 
contain free sulfur, this indicates that our organism belongs to the Sub-order 
Eubacteriineae. We note that the key to this suborder is on page 
67. 

We next attempt to ascertain the family to which the organism belongs 
by tracing it through the key to the familiesof the Sub-order Eubacteriineae, 
p. 67. 

I. No endospores indicates our organism. We proceed to A. Can develop 
on inorganic media. As the organism cannot grow without organic carbon, 
we turn to B. Cannot develop on inorganic media. 

This corresponds with the physiology of our organism; so we turn to 

1. Polar flagellate, etc. As our organism is peritrichous, we proceed to 

2. Large oval, pleomorphic cells sometimes almost yeast-like in appearance. 
Free living in soil. Fix free nitrogen. As this does not correspond with the 
morphology or physiology of our organism, we next examine 3. Peritrichous 
or non-motile rods, and cocci. This corresponds with the characteristics of 
our organism. 

We turn to a. Heterotrophicrodswhichmaynot require organic nitrogen for 
growth. Usually motile with one to six or moreflagella. Usually form nodules 
or tubercles on roots of plants, or shoio violet chromogenesis. 

This again does not indicate our oi canism ; so we turn next to aa. Heter- 
otrophic rods or cocci iDhich utilize organic nitrogen and usually carbohydrates. 
As our rod-shaped organism prefers a medium containing organic nitrogen, 
we proceed to b. Spherical cells in masses, tetrads and packets. 

This does not correspond to the morphology of our organism, and we now 
proceed to bb. Spherical cells ivhich grow in pairs and chains; and rods. 
This includes our rod-shaped organism; so we turn to c. Gram-positive 
cocci and rods. Non-motile. Since these are not the characteristics of our 
organism, we turn to cc. Gram-negative rods. When motile, from four to 
many peritrichous flagella. 

Our organism is Gram-negative and peritrichous; so we proceed to d. 
Grow well on ordinary media containing peptone. Aerobic to factdtative 
anaerobic. 

This corresponds with the characteristics of the organism we have 
studied; so we turn next to e. Gram-negative, straight rods which ferment 



SUGGESTIONS FOR USE OF MANUAL 3 

sugars with the formation of organic acids. This again corresponds with our 
organism. We turn next to f. Produce little or no acid from litmus milk. 
This does not correspond with the characters we have determined for our 
organism. We proceed to ff. Produce CO2 and frequently visible gas (CO2 + 
H2) from glucose. Reduce nitrates, etc. 

Our organism produces visible gas from glucose and reduces nitrates. 
This indicates that it belongs to Family X. Enterohacteriaceae, p. 443. 

This appears to fit our unknown organism. We now refer to page 443 
on which the key to the Family Enterohacteriaceae is found. In this key we 
ascertain the Tribe to which our organism belongs. 1. Ferment glu- 
cose and lactose with the formation of acid and visible gas. Usually do 
not liquefy gelatin. Tribe I. Eschericheae. 

This corresponds with the characters exhibited by our organism. We 
refer to the key for Tribe I. Eschericheae on the same page. 1. Methyl 
red test positive. Voges-Proskauer test negative. Salts of citric acid may or 
may not be used as sole source of carbon. Genus I. Escherichia, p. 444. 

This description appears to correspond with that of our unknown or- 
ganism. We find the key to the species of Genus Escherichia follows the 
key to the Tribe Eschericheae. On tracing our organism in this key we find 
that it corresponds to Escherichia coli. A brief description of this or- 
ganism is found on the same page. 

* In the use of keys for identifying bacteria, the student is confronted 
with two difficulties, both based primarily on lack of knowledge and ex- 
perience. The first is insufficient knowledge concerning the morphology, 
physiology, possible pathogenicity and habitat of the microorganisms that 
are to be identified. This may be due to careless observations or to poor 
training in the special techniques that must be used in determining the 
identity of a given bacterium. 

The second difficulty in the use of a key comes from inexperience in the 
use of technical terms; that is, the student may not thoroughly understand 
the meaning of the statement in the key and so cannot follow a route 
through the key with certainty. For example in the keys used here, the 
student must know the difference (1) between chains of cells which are 
composed of dividing cells which do not separate at once, and (2) filaments 
which are composed of dividing cells which remain more permanently to- 
gether and are normally flattened against each other on adjacent sides. 
They may show some differentiation into hold fast cells and reproductive 
cells (conidia), (3) Both chains of cells and filaments are to be distinguished 
from the mycelial threads found in Actinomycetaceae. These are unseptate 
and branching with a true branching. 

* Condensed and paraphrased from Hitchcock's Descriptive Systematic Botanj^, 
New York, 1935. 



4 SUGGESTIONS FOR USE OF MANUAL 

The student should be warned not to take descriptions in the Manual 
too literally or too rigidly. Descriptions are usually drawn to represent 
average findings. Especially among bacteria, characters such as sugar 
fermentations, gelatin liquefaction, presence or absence of flagella and 
other things will vary. Sometimes these variations are due to slight, 
possibly unrecognized variations in the techniques used in determining 
these characters. Real knowledge of the characteristics of species may also 
be very incomplete. This is true not only of the physiological activities 
of these microorganisms; but also in regard to such detectable structural 
features as the number and position of flagella. Dark field movies of motile 
cells and photographs taken with the recently developed electron micro- 
scope are revealing new and heretofore unsuspected facts regarding struc- 
tural features. 

Source and habitat data are frequently helpful in aiding the student 
to recognize species of bacteria and may indicate that the pathogenicity of 
the culture in question may need to be tried on some specific animal or 
plant. By habitat is meant the kind of a place in which the organism 
normally grows; by source, the particular material and place from which 
the culture was obtained. This source may or may not indicate the 
natural habitat. The source of cultures is invariably more limited in scope 
than the habitat as bacteria normally occur wherever their particular 
habitat may be found in a world wide distribution. 

The student is also reminded that it is impracticable to note all exceptions 
in keys. Bacteria like other living things are classified according to a 
combination of characters, not according to some single character, and ex- 
ceptions to the characters noted in the keys will occur in nature. These 
may not be known to or may have been overlooked by the author of the key. 
On the other hand, the importance of such exceptions should not be over- 
emphasized and the student would do well to use the key as if there were no 
exceptions. 



HISTORICAL SURVEY OF CLASSIFICATIONS OF 

BACTERIA, WITH E^IPHASIS ON OUTLINES 

PROPOSED SINCE 1923* 

There have been numerous attempts to arrange the species of bacteria in 
natural systems of classification. The first simple system of JMiiller (Ver- 
mium terrestrium et fiuviatilium, 1773) which he developed further a few- 
years later (Animalcula infusoria fluviatilia et marina, 1786) listed but two 
genera {Vibrio and Monas) that included organisms that would today 
probably be accepted as bacteria. Polyangium Link (^lag. d. Ges. Natur- 
forsch. Freunde zu Berlin, 3, 1809, 42) is apparently the oldest of the generic 
terms retained in its original meaning for a bacterial genus while Serratia 
Bizio (Biblioteca italiana o sia giornale de lettera, scienze ed arti, SO, 1823, 
288) was proposed only fourteen years later. 

Systems of classification developed after 1773 are given in complete 
outline form in the first edition of the ^Manual (1923) and this section of 
the IManual was reprinted without material change in the second (1925) 
and third (1930) editions. While it is not felt to be necessary to repeat 
these outlines in their entirety, sufficient reference is made below to permit 
the student to trace the origin of generic terms that are no longer commonly 
found in classification outlines. No attempt has been made to include 
reference to other little used generic terms except as the}- appear as syn- 
onyms in the descriptive portion of the ^Manual. For the origin of generic 
terms proposed before 1925, see Enlows (The Generic Names of Bacteria, 
Bui. No. 121, Hygienic Laborator}-, Washington, D. C, 1920) and Bu- 
chanan (General Sj-stematic Bacteriology, Baltimore, 1925). 

Bory St. Vincent (IMicroscopiques, Dictionnaire classique d'histoire 
naturelle, 10, 1826, 533) introduced the generic terms Spirilina, Melanella, 
Lactrimatoria and Pupella and accepted Vibrio for microorganisms, some of 
which must have been bacteria. None of these terms, except Vibrio, are in 
current use for bacterial groups. 

Three of the terms accepted or proposed by Ehrenberg (Die Infusions- 
tierchen als volkommene Organismen, Leipzig, 1838); nameh^, Vibrio, 
Spirillum and Spirochaeta, are still used. The generic term Bacterium 
proposed first b}- Ehrenberg in 1828 (SjTiibolae Physicae seu Icones et 
Descriptiones Animalium Evertebratorum Separasitis Insectis quae ex 
Itinere per Africam Borealem et Asiam Occidentalem, IV. Evertebrata, 
Berlin) to include but a single species Bacterium triloculare from an oasis 

* Contributed by Prof. R. S. Breed, New York State Experiment Station, Geneva, 
New York, July, 1938; revised, September, 1943. 



b MANUAL OF DETERMINATIVE BACTERIOLOGY 

in North Africa, has had a varied history because this type species (mono- 
typy) is no longer identifiable. It was reintroduced into the classification 
employed in the fifth edition of the Manual to cover species of non-spore- 
forming rods whose positions in the outline given in the Manual have not 
yet been satisfactorily determined (Breed and Conn, Jour. Bact., 31, 1936, 
517) and is used in the present edition with the same meaning. The term 
Spirodiscus was applied by Ehrenberg to a single organism that he found in 
a mountain stream. It has never been reidentified and subsequent authors 
have discarded this term. 

Two new generic terms {Metallacter, Sporonema) were introduced by 
Perty (Zur Kenntniss kleinster Lebensformen, 1852). Neither Metallacter 
nor Sporonema is in common use at the present time. 

Davaine (Dictionaire encyclop. des sciences med., Art. bacteries, 1868) 
introduced one new generic term, Bacteridium, for straight motionless rods 
like the anthrax bacillus. 

The generic terms employed by Cohn in his first classification (Unter- 
suchungen iiber Bakterien. I. Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 
1872, 146) are all in current use. Only one {Bacillus) was new. Other 
generic terms were introduced into his second paper (Untersuchungen tiber 
Bakterien. II. ibid., 1, Heft 3, 1875, 141) which contained his more 
complete classification. For various reasons, six of these, Merismopedia, 
Clathrocystis, Ascococcus, Myconostoc, Cladothrix and Streptothrix are not 
found in recent bacteriological classifications. 

Mangin (Les Bacteries, Paris, 1878) recognized three subgenera of the 
genus Monas, the first of which Rhabdomonas Cohn, 1875 is still used as a 
generic term, while the other two, Ophidomonas Ehrenberg, 1838 and Spiro- 
monas Perty, 1852 have been dropped. 

The bacterial species that had been placed in the genus Clathrocystis by 
Cohn (1875) were separated and placed in a new genus Cohnia by Winter 
(Die Pilze in Rabenhorst's Kryptogamen Flora, 1880), and this name is also 
used by Burrill (The Bacteria, Springfield, 111., 1882). Because this name 
had previously been proposed for a genus of lilies, it was soon dropped. 

Zopf (Die Spaltpilze, Leipzig, 1883) accepts Phragmidioihrix, a generic 
name suggested by Engler in 1882 for a single species found on the body of 
a crustacean {Gammarus locusta). Later authors generally either merge 
this genus with Crenothrix Cohn or disregard it because of the indefinite 
description of the one species included in it. 

Baumgarten (Lehrbuch der pathologischen Mykologie, Braunschweig, 
1890) following Hueppe accepts the term, Spirulina, for a genus of pleo- 
morphic bacteria, disregarding the previous use of the term by algologists. 

The generic terms found in Migula's first outline (Bakterienkunde fiir 



SURVEY OF CLASSIFICATIONS OP BACTERIA 7 

Landwirte, Berlin, 1890) were those in conventional use at the time and 
many of them continue in use. Two new terms were introduced for motile 
types in his second outline (Ai'b. Bact. Inst. Karlsruhe, 1, 1894, 235) and 
are also found in his later outlines (Engler and Prantl, Die natiirlichen 
Pflanzenfamilien, 1, la, 1895, 29, and System der Bakterien, 1, 1897, 46, 
and 2, 1900, 269 and 275) which have not been generally felt to be necessary 
by subsequent authors. These are Planococcus and Planosarcina. Spiro- 
soma introduced by Migula in 1894 and Rhabochromatium Winogradsky ac- 
cepted by Migula in 1900 are like^\ise no longer generally used. Newskia 
(original spelling Nevskia Famintzen, Bull. Acad. Imp. Sci., St. Petersburg, 
34 (N.S. 2), 1892, 484) has recently been revived by Henrici and Johnson 
(Jour. Bact., 29, 1935, 3 and 30, 1935, 83). The generic term Microspira 
Schroeter, accepted by Migula in 1894, is still frequently accepted in place 
of Vibrio as many regard it as having a better status than the later term. 

The term Pseudomonas was first proposed for polar flagellate bacteria by 
Migula in his 1894 outline with reference to but a single species, Pseudo- 
monas violacea, an organism which later investigators have shown to be 
peritrichous (Cruess-Callaghan and Gorman, Sci. Proc. Roy. Dublin Soc, 
21, 1935, 213). Pseudomonas was repeated in the 1895 outline with 
descriptions of Pseudomonas pyocyanea and other species. Later authors 
have generally accepted the term Pseudomonas as valid. 

Fischer (Jahrb. f. mssensch. Bot., Berlin, 27, 1895, 1) introduced a 
logical outline classification in which he proposed various new terms which 
have never come into general use. These are Paracloster, Paraplectrum, 
Arthrobader, Bactrinium, Clostrinium, Plectrinium, Arthrobactrinium, 
Bactrillum, Clostrillium, Plectrillium, Arthrobactrillium, Badridium, Plec- 
tridium, Dipledridium, and Arthrobadridium. In his modified classifica- 
tion (Vorlesungen iiber Bakterien, 1897), he also accepts Pediococcus 
Balcke, a term that has fallen into disuse except in the brewing indus- 
try. 

In the conservative classification proposed bj'' Lehmann and Neumann 
(Atlas und Grundriss der Bakteriologie, 2 vols., 1896, Miinchen), inter- 
nationally accepted rules of nomenclature were followed. All of the generic 
terms employed by them are still in current use, their most important 
contribution being their acceptance of the suggestion that the genus 
Bacillus be separated from the genus Baderium on the basis of endospore 
formation by the rods included in Bacillus. Two new genera were proposed 
{Corynebaderium and Mycobacterium) that have been generally accepted 
by later workers. 

No new generic terms are proposed by Chester either in his preliminarj'^ 
reports (Delaware College of Agriculture, 9th Ann. Kept., 1897, 53 and 62; 



8 MANUAL OF DETERMINATIVE BACTERIOLOGY 

11th Ann. Rept., 1899, 36), or in his complete outhne (Manual Determ. 
Bact., 1901). Almost all of the generic terms found in his outlines are still 
in current use. 

The term Aplanohacter suggested by Erwin F. Smith (Bacteria in Rela- 
tion to Plant Diseases, 1, 1905, 171, Washington) was accepted by certain 
American phytopathologists for a time but has never come into general use. 

Because other differences between the non-chromogenic and chromogenic 
micrococci are unimportant, two generic terms, Albococcus and Aurococcus, 
suggested by the Winslows (Science, 21, 1905, 669; Systematic Relation- 
ships of the Coccaceae, New York, 1908) have not come into general use. 
They also suggested Rhodococcus to include Rhodococcus roseus and R.fulvus 
apparently without realizing that Zopf (Ber. d. deutsch. bot. Gesellsch. 
Berlin, 9, 1891, 28) had previously used the same term for Rhodococcus 
erythromyxa and R. rhodochrous. Hansgirg (Engler and Prantl, Die 
natiirhchen Pflanzenfamilien, 1, la, 1895, 52) had also used it previously 
to designate a sub-genus of the green algae, and later Alolisch (Die Purpur- 
bakterien, Jena, 1907, 20) used Rhodococcus for a genus of the purple 
bacteria to include Rhodococcus capsulatus. 

In his complete outline of the classification of bacteria presented in 1909, 
Orla-Jensen (Cent. f. Bakt., II Abt., 22, 1909, 305) introduced many new 
generic terms in an effort to create a nomenclature that appeared to him to 
express the natural relationships of bacteria more satisfactorily than names 
previously suggested had done. Thus he used the suffixes coccus and 
sarcina for spherical bacteria and monas for all genera known to be lopho- 
trichous or so related to these types that they were regarded as essentially 
lophotrichous in nature. In the same way the suffix bacterium was used 
for genera of non-spore-forming rods that were regarded as essentially 
peritrichous in nature, and the suffix bacillus for similar spore-forming rods. 
As, however, subsequent investigators have (1) accepted the priority rule, 
(2) felt that it was impossible to recognize the type of motihty found in the 
ancestry of truly non-motile groups, or (3) felt that other characters were 
more fundamental than those selected by Orla-Jensen, many of these terms 
have not been generally used by later workers. 

Among the little used terms suggested or accepted by Orla-Jensen are: 
Acetimonas, Nitromonas, Azotomonas, Rhizomonas, Corynemonas, Myco- 
monas, Sulfomonas, Thiomonas, Thiococcus, Rhodomonas, Rhododictyon, 
Amoebomonas, Rhodopolycoccus, Rhodosarcina, Spirophyllum, Denitro- 
monas, Liquidomonas, Liquidovibrio , Liquidococcus, Solidococcus, Solido- 
vibrio, Sporosarcina, Denitrobacterium, Caseobacterium, Liquidobacterium, 
Urobacillus, Butyribacillus, Pectobacillus, Cellulobacillus, Putribacillus and 
Botulobacillus. 



SURVEY OF CLASSIFICATIONS OF BACTERIA 9 

While Nitromonas is not new, it is redefined as a synonym of Nitrohader 
Winogradsky (Arch. Sci. Biol. St. Petersburg, 1, 1892, 87), rather than as a 
synonym of Nitromonas Winogradsky (Ann. Inst. Past., 3, 1890, 258). 
Spirophyllum is from Ellis (Cent. f. Bakt., II Abt., 19, 1907, 507). 

In a later monograph on The Lactic Acid Bacteria (Mem. d. Acad. Roy. 
Sci. et Lettres de Danemark, Sect. Sci., 8 Ser., 5, 1919, No. 2) Orla-Jensen 
proposes the following additional generic terms: Betacocciis,Betahacteriuni, 
Streptobacterium, Thermohacteriiim and Microhacterium. The term Tctra- 
coccus is introduced with a meaning different from that given the term 
previously by v. Klecki (Cent. f. Bakt., 15, 1894, 354). 

Buchanan prepared an outline classification in 1916 (Jour. Bact., 1, 
1916, 591 ; 2, 1917, 155, 347, 603; 3, 1918, 27, 175, 301, 403, 461, 591) which 
was utilized in part by the group of which he was a member (Winslow, 
Broadhurst, Buchanan, Krumwiede and Smith) in their preliminary Report 
to the Society of American Bacteriologists (Jour. Bact., £, 1917, 552) and 
in the final report by Winslow, Broadhurst, Buchanan, Krumwiede, Rogers 
and Smith (Jour. Bact., 5, 1920, 191). 

Although prepared earlier, some parts of the Buchanan outline were not 
published until after the first Winslow et al. report. As these reports 
formed the most important basis for the classification used in the first edi- 
tion of the Manual, it is natural that the generic terms utilized are, in 
general, the same as those used in the Manual. 

Generic and subgeneric terms included by Buchanan that are not used 
in the present edition of the Manual are: Paraspirillum Dobell (Arch. f. 
Protistenk., 21^, 1911, 97), Eubacillus Hansgirg (Osterr. Bot. Ztschr., 38, 
1888, 264; not Eubacillus Dangeard, Le Botaniste, 2, 1891, 151) and Meta- 
bacterium Chatton and Perard (Comp. rend. Soc. Biol., Paris, 65, 1913, 
1232). Siderocapsa Molisch (Ann. Jard. Bot. Buitenzorg, Ser. 2, Supp. 
3, 1909, 29) used by Buchanan but dropped by Winslow et al (Jour. Bact., 
2, 1917, 549) does not appear in the Manual classification outline until the 
present (6th) edition. The term Mycoderma recognized both })y Buchanan 
(Jour. Bact., 3, 1918, 45) and in the preliminary Winslow et al. report 
(Jour. Bact., 2, 1917, 551) was replaced by the later and more valid term 
Acetobacter in the final report by WinsloAv et al. (Jour. Bact., 5, 1920, 201). 
Pfeifferella Buchanan (Jour. Bact., 3, 1918, 54) which is used in the three 
outline classifications under discussion and also in the first, second and third 
editions of the ^Ianual, appeared in the literature through a clerical error 
(Buchanan, General Systematic Bacteriology, 1925, 420) . It was combined 
in the fourth edition of the Manual with the genus Actinobacillns under the 
latter name. Nocardia Trevisan (1889) used by Buchanan and in the 
preliminary report by Winslow et al. (1917) was merged with Actinomyces 



10 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Harz (Jahresber. Miinchen. Thierarzneisch. for 1877-78, 125) in the final 
report by Winslow et al. Erythrohacillus Fortineau (Compt. rend. Soc. Biol. 
Paris, 58, 1905, 104) is used by Winslow et al. (1920) but was not accepted 
in the first and following editions of the IVIanual as it is a synonym of the 
older Serratia Bizio (1823). Moreover, the species which must be accepted 
as type for the genus {Erythrohacillus pyosepticus Fortineau (monotypy)) 
is a species which has been reported by Breed (Manual, 3rd ed., 1930, 117) 
to be a variant of the older Serratia marcescens. 

One of the most unsatisfactory portions of recent classifications, such as 
those outlined by Buchanan (1917-18) and by Winslow et al. (1917), is 
the treatment given the organisms of the coliform-dysentery-typhoid group 
in that the term Bacterium is retained for these as suggested by Orla- 
Jensen (1909). A strict limitation of Bacterium to this group gives it a still 
different meaning from that which it had had in previous and current 
classifications, and makes it necessary to find some other place for many 
other species of Gram-negative, non-spore-forming rods, some of which are 
well known and well described. The relationships of these miscellaneous 
species to other non-spore-forming rods is frequently poorly understood. 
In some cases, further study will probably show that thej'' should be placed 
in well known and currently recognized genera. In others, further study 
will probably show that some of these species of non-spore-forming rods 
should be grouped in new genera. 

Winslow et al. (1920) recognized this situation and broadened their defini- 
tion of Bacterium thereby placing such w^ll known species as are included 
in the colon-dysentery-typhoid group with other species of non-spore-form- 
ing rods of quite a different character. For this reason, partial use was 
made in the first edition of the Manual of the numerous generic terms newly 
proposed by Castellani and Chalmers (Manual of Tropical Medicine, 3rd 
ed., 1919). Thus the following new terms were introduced: Alcaligenes, 
Salmonella, Escherichia and Encapsulatus; and the earlier terais Aerohacter 
Beijerinck (1900) and Eberthella Buchanan (1918). Later it was found that 
Encapsulatus was a synon3^m of Klebsiella Trevisan (1887), so that the latter 
term was accepted in the second and subsequent editions of the Manual. 
Shigella Castellani and Chalmers was recognized as distinct from Eberthella 
in the third and subsequent editions. 

Many of the new terms suggested by Castellani and Chalmers were, 
however, synonyms of earlier valid terms or have not been considered 
necessary, and so they have not come into general use. These are Nigro- 
coccus, Graciloides, Cloaca, Eberthus, Dysenteroides, Lankoides, Wesenbergus, 
Balkanella and Enter oides. No new generic terms are given by Castellani 
and Chalmers in their later report (Ann. Inst. Past., 34, 1920, 600). 



SURVEY OF CLASSIFICATIONS OF BACTERIA 11 

Orla-Jensen (Jour. Bact., 6, 1921, 263), in a paper published after the 
manuscript of the first edition of the Manual was prepared, suggested the 
use of Colibacterium and Aerogeneshacterium for the two genera in the 
coliform group and adds quite a number of other new terms formed in 
accordance with his system of nomenclature. These are, in most cases, 
synonjTns of earlier valid names. The new terms are Coccomonas, Spiro- 
monas (used in a new, different sense from that of earlier authors), Fluoro- 
monas, Photomonas, Propionicoccus, Buty rich sir idium and Putriclostridium. 

Many new terms are proposed in the classification drawn up by Heller 
(Jour. Bact., 6, 1921, 521; and 7, 1922, 1). Details are given in the group 
of anaerobic spore-formers onlj*. Here each of the new generic terms is 
based on a single species. The following outline is given in the first of these 
papers, two new genera {Rivoltillus and Metchnikovillus) being made the 
type genera for two new subfamilies Clostridioideae and Putrificoideae, 
respectively. 

Phylum I. Bacteria 

Class I. Eubacterieae 

Order 1. Eubacieriales 

Family C (?). Clostridiaceae 

Subfamily 1. Clostridioideae 
Subfamily 2. Putrificoideae 
Order 2. Thiobacteriales 
Order 3. Chlamydohacteriales 
Class II. Myxohacterieae 

In the more complete outline in the second paper, one generic term 
(Clostridium) is old, although it is used in a new and restricted sense, while 
with the exception of the type genera mentioned above, the other terms 
are new. In the subfamily Clostridioideae, the new terms are Omelianskillus, 
Macintoshillus, Douglasillus, Henrilliis, Flemingillus, Vallorillus, Multi- 
fermentans, Hihlerillus, Welchillus, Stoddardillus, Arloingillus, Meyerillus 
and Novillus. Ten new generic terms are used in the subfamily Putrifi- 
coideae as follows: Sequinillus, Reglillus, Rohertsonillus, Nicollaierillus, 
Martellillus, Recordillus, Tissierillus, Putrificus, Ermengemillus, and Wein- 
bergillus. As there does not seem to be any good reason for sub-dividing 
the genus Clostridium in this way, the latter term has been used to cover 
anaerobic spore-forming rods in all previous editions of the Manual, and is 
again used in the present edition in this sense rather than with the restricted 
meaning proposed bj^ Heller. 

Enderlein (Sitzber. Gesell. Naturf . Freunde, Berlin, 1917, 309) proposed 
an outline classification covering the Kingdom of Mychota, or bacteria, 
which was based on comparative morphology with special emphasis on life 
cycles. This was as follows: 



12 MANUAL OF DETERMINATIVE BACTERIOLOGY 



Phylum I. Dimychota 


Ki 


reis A. Holocyclomorpha 




Class I. Gonascota 




Order a. Synascota 




Family 1. Schaudinnidae 




Genus a. Schaudinnum 




b. Theciobactrum 




Family 2. Sphaerotilidae 




Genus a. Phragmidiothrix 




b. Newskia 




c. Chlaynydothrix 




d. Sphaerotilus 




e. Clonothrix 




Family 3. Syncrotidae 




Genus a. Crenolhrix 




h. Beggiatoa 




c. Syncroiis 




d. Zygoslasis 




Family 4. Spirillidae 




Genus a. Gallionella 




b. Spirillum 




c. Dicrospirillum 




Family 5. Spirochaeiidae 




Genus a. Cristispira 




h. Treponema 




c. Entomospira 




d. Spirochaeta 




e. Cacospira 




Family 6. Microspiridae 




Genus a. Spirobacillus 




b. Spirosoma 




c. Photobacterium 




d. Microspira 




e. Dicrospira 




Family 7. Corynobacteriidae 




Subfamily 1. Aclinomycinae 




Genus a. Actinomyces 




Subfamily 2. Eisenberginae 




Genus a. Eisenbergia 




Subfamily 3. Sclerolrichinae 




Genus a. Zetlnowia 




b. Schlerothrix 




Subfamily 4. Corynobacteriinae 




Genus a. Corynobaclerium 




h. Heterocyslia 




c. Cladascus 




d. Zygoplagia 




Subfamily 5. Pseudosirepinae 




Genus a. Pseudostreplus 




Order b. Ascoia 




Family 8. Bacteriidae 



SURVEY OF CLASSIFICATIONS OF BACTERIA 13 

Genus a. Atremis 

b. Bacterium 

c. Lamprella 

d. Eucystia 

e. Dicrobactrum 

f. Acysiia 
Family 9. Fusiformidae 

Genus a. Fusiformis 
Class II. Sporascota 

Order a. Parasynascota 
Family 10. Migulanidae 

Genus a. Migulanum 
Order b. Parascota 

Family 11. Bacillidae 

Genus a. Rhagadascia 
h. Plectridium 

c. Bacillus 

d. Bacirillum 

e. Kochella 

f. Fischerinum 
Kreis B. Hemicyclomorpha 

Class I. Anascola 

Familj' 12. Hemallosidae 
Genus a. Hemallosis 
Phylum II. Monomychota 
Kreis A. Acyclomorpha 

Family 1. Mogallidae 

Genus a. Mogallia 
Family 2. Sarcinidae 

Genus a. Diplococcus 

b. Sarcina 

c. Paulosarcina 
Family 3. Micrococcidae 

Genus a. Micrococcus 

h. Planococcus 

c. Streptococcus 

d. Phacelium 

Three of the new generic term.s, Cladasciis tj^pe species C. furcabilis 
Enderlein, Zygoplagia type species Z. alternans Enderlein and Heterocystia 
type species H. multiformiis Enderlein, had been proposed in an earHer 
paper (Sitzber. Gesell. Naturf. Freunde, BerUn, 1916, 395). The following 
generic terms in the 1917 outline are new: Schaudinnum, TJieciobactrum, 
Syncrotis, Zygostasis, Dicro spirillum, Entomospira, Cacospira, Dicrospira, 
Eisenhergia, Zettnoivia, Pseudostreptus, Atremis, Lamprella, Eucystia, Dicro- 
bactrum, Acystia, Migulanum, Rhagadascia, Kochella, Fischerinum, Hemal- 
losis, Mogallia, Paulosarcina and Phacelium. Note that Corynobacterium 
is spelled with an o instead of an e. 

Terms accepted from earlier workers that have not previously been 



14 MANUAL OF DETERMINATIVE BACTERIOLOGY 

mentioned are: Spirobacillus Metschnikoff (Ann. Inst. Past., 3, 1889, 62), 
Photohacterium Beijerinck, Maanblad voor Natuurwetenschappen Amster- 
dam, 16, 1889, 1 and Arch. Ne^rl. d. Sci. Exactes, 23, 1889, 401), and 
Sclerothrix Metschnikoff (Arch. f. Path. Anat. u. Physiol., 113, 1888, 
63-94; not Sclerothrix Kuetzing, Species Algarum, 1849, 319). 

The above outHne was changed in 1925, p. 235 ff. (Bakterien-Cyclogenie, 
BerHn, 390 pp.) by the addition of one new family, Chondromycidae, to 
include the genus Newskia, formerly included in Sphaerotilidae, and nine 
genera not previously given as follows : Chondromyces, Cystodesmia, Mono- 
cystia, Ophiocystia, Apelmocoena, Polyangium, Cystoecemia, Myxococcus and 
Dactylocoena. All except Chondromyces, Polyangium and Myxococcus are 
taken from Enderlein (Bemerkungen zur Systematik der Chondromyciden, 
Berlin, 1924, 6 pp.). 

The new genus Lohnisium is added in the Family Eisenhergiinae to in- 
clude the acetic acid and legume bacteria, and he also proposes the generic 
term Macrocystita (p. 278) for certain bacteria described by Peklo (0 
msici krvave (Study of the blood louse). Zemedelskdho Archivu (Agri- 
cultural Archives), 1, 1916) from aphids. According to Enderlein it is not 
clear whether this genus should be included in the Family Bacteriidae or in 
Corynobacteriidae . 

Two genera proposed by others are also accepted. These are Calym- 
matohacterium Aragao and Vianna (Mem. Inst. Oswaldo Cruz, 6, 1912, 211) 
placed in the family Migulanidae, and Leuconostoc Van Tieghem placed in 
the family Micrococcidae. 

Later Enderlein (Sitzber. Gesell. Naturf. Freunde Berlin, 1930, 104-105) 
accepts Serratia Bizio in place of Dicrohactrum, and Leptotrichia Trevisan in 
place of Syncrotis. Streptus with Streptus scarlaiinae as type species, is 
proposed to cover the streptococci not included in Pseudostreptus. 

The outline suggested by Pringsheim (Lotos, 71, 1923, 357) is similar to 
that used by Lehmann and Neumann (Atlas und Grundriss der Bakterio- 
logie, 2 vols., 1896, Miinchen). It is a conventional division into spherical, 
rod-shaped and curved forms so far as the true bacteria are concerned except 
that the pseudomonads are included in the same family as the vibrios and 
spirilla. Rhodohacteriales is recognized as an order to include the sulfur 
purple bacteria and the nonsulfur purple bacteria. Few details are given 
in regard to the other orders. His outline follows: 

Schizomycetes 

Order I. Eubacteriales 
Family 1. Coccaceae 

Genus a. Streptococcus 

b. Micrococcus 

c. Sarcina 



SURVEY OF CLASSIFICATIONS OF BACTERIA 15 

Family 2. Bacleriaceae 

Genus a. Bacterium 
b. Bacillus 
Family 3. Spirillaceae 

Genus a. Pseudomonas 

b. Vibrio 

c . Spirillum 
Order II. Rhodobacteriales 

Family 1. Rhodobacterinae 
2. Thiorhodinae 
Order III. M yxobacteriales 

Family 1. Myxobacteriaceae 
Order IV. Mycobacteriales 

Family 1. Corynebacteriaceae 

2. Mycobacteriaceac 

3. Actinomycelaceae 

(Also possibly the long rod, lactic acid bacteria.) 
Order V. Desmobacteriales 

Family 1. C hlamydobacteriaceae 
2. Beggiatoaceae 

The first outline classification drawn up by Janke (Allgemeine Technische 
Mikrobiologie, I Teil, Dresden, 1924, p. 63) is an adaptation and expansion 
of that drawn up by Migula (System der Bakterien, 1900) . The new genera 
recognized by Janke are Planostre^tococcus A. Meyer (Die Zelle der Bak- 
terien, Jena, 1912), Thioploca Lauterborn (Ber. dtsch. Bot. Gesell,, 25, 
1907, 238), Thiohacterium Molisch (Cent. f. Bakt., II Abt., 33, 1912, 55), 
Thiobacillus Beijerinck (Cent. f. Bakt., II Abt., 11, 1904, 593), Thiovihrio 
Janke {loc. cit.), Thiospirillum Winogradsky (Beitrage zu Morphol. u. 
Physiol, d. Bakterien. Heft I. Schwefelbakterien. Leipzig, 1888), Thio- 
sphaerella Nadson (Bull. Jar. bot. Petersburg, 13, 1913, 106; ref. in Cent. 
f. Bakt., II Abt., 43, 1915, 469), Thiovulum Hintze (Ber. Dtsch. Bot. 
Gesell., 31, 1913, 189), Spirophyllum Ellis (Proc. Roy. Soc. Edinburgh, 27, 
I, 1907, 21; ref. in Cent. f. Bakt., II Abt., 19, 1907, 502), Nodofolium Ellis 
(Cent. f. Bakt., II Abt., 26, 1910, 321), and Actinococcus Beijerinck (Fol. 
Microbiol.,:^, 1914, 185). 

Janke 's outline classification is given below: 

Order I. Eubacteria 
Family 1. Coccaceae 

Genus a. Streptococcus 
h. Micrococcus 

c. Sarcina 

d. Planostreptococcus 

e. Planococcus 

f. Planosarcina 
Family 2. Bacleriaceae 

Genus a. Bacillus 
b. Bacterium 



IQ RIANUAL OF DETERMINATIVE BACTERIOLOGY 

Family 3. Spirillaceae 

Genus a. Microspira 

b. Spirillum 

c. Spirosoma 
Order II. Rhodobacteria 

Family 1. Thiorhodaceae 
Subfamily la. Thiocysteae 
Genus a. Thiocystis 

b. Thiocapsa 

c. Thiosphaera 

d. Thiosphaerion 

e. Thiosarcina 
Subfamily 2b. Lamprocysteae 

Genus a. Lamprocystis 
Subfamily 3c. Thiopedieae 
Genus a. Thiopedia 
h. Thioderma 
Subfamily 4d. Amoebobacterieae 
Genus a. Amoebobacter 

b. Thiothece 

c. Thiodictyon 

d. Thiopolycoccus 
Subfamily 5e. Chromatieae 

Genus a. Chroviatium 

b. Rhabdochromatium 

c. Thiorhodospirillu7n 
Subfamily 6f. Rhodocapseae 

Genus a. Rhodocapsa 

b. Rhodothece 
Family 2. Athiorhodaceae 

Subfamily la. Rhodocysteae 

Genus a. Rhodocystis 

h. Rhodonostoc 

c. Rhodococcus 

d. Rhodobacterium 

e. Rhodobacillus 

f. Rhodovibrio 

g. Rhodospirillum 
Order III. Thiobacteria 

Family 1. Beggiatoaceae 
Genus a. Thiothrix 
h. Beggiatoa 
c. Thioploca 
Family 2. Thiobacteriaceae 
Genus a. Thiophysa 

h. Thiobacterium 

c. Thiobacillus 

d. Thiovibrio 

e. Thiospirillum 

f. Thiosphaerella 

g. Thiovulum 

h. Achromatium 



SURVEY OP CLASSIFICATIONS OP BACTERIA 17 

Order IV. Phycobacteria 

Genus a. Leptothrix 

b. Clonothrix 

c. Cladothrix 

d. Crenolhrix 

e. Phragmidioihrix 

Appendix Genera Gallionella, Spirophyllum, Nodofolium 
Order V. Mycobacteria 

Family 1. Mycobacteriaceae 

Genus a. Cory neb acterium 
b. Mycobacterium 
Family 2. Actinomycetaceae 
Genus a. Actinomyces 
b. Actinococcus 
Order VI. Myxobacteria 

Familj^ 1. Myxobacteriacene 
Genus a. Myxococcus 

b. Chondromyces 

c. Polyangium 

Lehmann and Neumann (Bakt. Diag., 2 vols., 7th ed., Miinchen, 1926-27 ; 
Breed, Eng. trans., New York, 1931) developed their first simple and much 
used outline classification, dra\vn up in 1896, in later editions of their 
Determinative Bacteriology. The 1927 Lehmann and Neumann outline 
is as follows: 

Class I. Schizomycetes 

Order I. Schizomycetales 
Family 1. Coccaceae 

Genus a. Streptococcus 

b. Sarcina * 

c. Micrococcus 
Sub-genus a. Diplococcus 

b. (Gram-positive group) 
Family 2. Bacleriaceae 
Genus a. Bacterium 

Sub-genus a. Nitrosomonas 

b. Nitrobacter 

c. Rliizobium 

d. Haemophilus 

e. Brucella 

f. Pasteurella 

g. (Glanders and dysenterj' group) 
h. (Photogenic group) 

i. (Aerogenes group) 

j. Encapsulatus 

k. (Typhoid group)* 

1. Salmonella 

m. (Coli group)* 

* In a footnote under these groups, the authors refer to the names given by Castel- 
lani and Chalmers. 



18 MANUAL OF DETERMINATIVE BACTERIOLOGY 





n. Aceiobacierium 




o. (Cloacae group) 




p. (Red chromogens) 




q. (Blue and violet chromogens) 




r. Pseudornonas 




s. Proteus 




App. Erysipelothrix 


Genus 


h. Fusobaclerium 




c. Plocamobacierium 


Family 3. 


Desmobacteriaceae 


Genus 


a. Beggiaioa 




h. Leplothrix 


Sub 


i-genus a. Leplothrix 




b. Chlamydothrix 


Genus 


c. Crenothrix 




d. Cladolhrix 




e. Thiothrix 


Family 4. 


Spirillaceae 


Genus 


a. Vibrio 




b. Spirillum 


Family 5. 


Spirochaetaceae 


Genus 


a. Spirochaela 


Family 6. 


Bacillaceae 


Genus 


a. Bacillus 


Sub 


i-genus a. (Aerobic group) 




b. (Anaerobic group) 


Order II. Actinomyceiales 


Family 1. 


Proactinomycelaceae 


Genus 


a. Corynebaclerium 




b. Mycobacterium 


Family 2. 


A ctinomycetaceae 


Genus 


a. Actinomyces 



The generic term Bacterium is retained in this outUne to cover those 
groups of the true bacteria that are Gram-negative, non-spore-forming, 
motile and non-motile rods. Lehmann and Neumann recognize 20 sub- 
groups in this genus, many of which correspond with the genera recognized 
in the Manual. In an effort to develop a rational nomenclature the term 
Acetobacterium (occurs first in review by Ludwig, Cent. f. Bakt., II Abt., 
4, 1898, 870) is used in place of Acetohacter, Plocamohacterium (Loewi, Wien. 
klin. Wchnschr., 33, 1920, 730) in place of Lactohacillus, and Fusobacterium 
(Knorr, Cent. f. Bakt., I Abt., Orig., 89, 1922, 4) in place of Fusiformis 
without regard to priority. Encapsulatus Castellani and Chalmers (Man- 
ual Tropical Med., 3rd ed., 1919, 934) is used in place of Klebsiella Trevisan 
(Atti Accad. Fis.-Med.-Stat. Milano, Ser. 4, 3, 1885, 107). 

Janke (Cent. f. Bakt., II Abt., 66, 1926, 481) reprints the classification 
developed in the first edition of the present Manual and compares it with 
that proposed by Orla- Jensen and Enderlein. 



SURVEY OF CLASSIFICATIONS OF BACTERIA 19 

The second complete outline drawn up by Janke (Oesterr. Bot. Zeitschr., 
78, 1929, 108) is similar to the classification employed by Lehmann and 
Xeumann (Bakt. Diag., 2 vols., 7th ed., Miinchen, 1926-27). He follows 
Enderlein in placing Azotohacter in close association with the spore-forming 
rods. No new generic terms are suggested. His sub-groups of the genus 
Bacterium are even more closely similar to the genera used in the present 
edition of the jNIaxual than are the sub-groups of Lehmann and Neumann. 

Family 1. Coccaceae 

Genus a. Micrococcus 

b. Neisseria 

c. Streptococcus 

Divided into 4 groups. 

d. Sarcina 

Divided into 2 groups. 
Family 2. Bacillaceae 
Genus a. Bacillus 

Divided into 16 groups, 
b. Azotohacter 
Family 3. Bacteriaceae 
Genus a. Bacterium 

Divided into 27 groups. 

b. Fusifortnis 
Family 4. Corynobacteriaceae 

Genus a. M ycohacterium 
h. Corynobacierium 

c. Actinomyces 
Family 5. Spirillaceae 

Genus a. Microspira 

Divided into 2 groups, 
b. Spirillum 

Divided into 2 groups. 



Family 6. 


Spirochaetaceae 


Genus 


a. 


Spirochaeta 




b. 


Borrelia 




c. 


Treponema 




d. 


Cristispira 




e. 


Saprospira 




f. 


Leptospira 


Family 7. 


Desmobacleriaceae 


Genus 


1 a. 


Beggiatoa 




b. 


Thioploca 




c. 


Thiothrix 




d. 


Leptolrichia 




e. 


Crenothrix 




f. 


Sphaerotilus 




S- 


Clonothrix 




h. 
i. 


Leptothrix 
Phragmidiothrix 



20 ' MANUAL OF DETERMINATIVE BACTERIOLOGY 

Family 8. M yxobacteriaceae 
Genus a. Myxococcus 

b. Polyangium 

c. Chondromyces 

Pribram (Jour. Bact., 18, 1929, 361) has rearranged some groups and 
combined others (e.g., Rhizohium, Diplococcus, Leuconostoc, Serratia, 
Flavohacterium, Chromobacterium, Achromohader, Cellulomonas) recognized 
in the first edition of the Manual with Httle change in the nomenclature 
except among the anaerobic non-spore-forming rods and among the spore- 
forming rods. Unfortunately, he has sometimes used family and species 
names as generic names, thus in the latter case introducing adjectives and 
adjectival terms as substantives. New generic terms suggested are: 
Dialisterea, Bacteroidea, Centrosporus, Fusibacillus, Pseudobacillus, Mega- 
therium, Flexus, Anthrax, Botulinus, Chauvoea, Botidinea, Putrificus, 
Welchia, Phleobacterium, Distasoa, Tissieria, and Actinoidomyces. Astasia 
as it appears in this outline does not appear to be the same as Astasia Meyer 
(Flora, 84, 1897, 185). Aerobacillus is not synonymous with Aerobacillus 
Donker (Inaug. Diss., Delft, 1926). Sideromonas is accepted from Cho- 
lodny (Ber. Deutsch. Bot. Ges., 40, 1922, 326). 

Pribram's complete outline follows: 

Class Schizomycetes 

Subclass A. Prolozoobacteria 
Order I. Spirochaetales 

Family 1. Spirochaetaceae 
Genus a. Spirochaeta 

b. Treponema 

c. Spironema 
Family 2. Crisiispiraceae 

Genus a. Saprospira 
h. Cristispira 
c. Leptospira 
Subclass B. Eubacleria 
Order I. Protobacteriales 

Family 1. Nitrobacteriaceae 
Related to Pseudomonas 
Tribe A. H ydrogenomonadae 
Genus a. H ydrogenomonas 

b. M ethanomonas 

c. Carboxyodomonas 
Tribe B. Niirobactereae 

Genus a. Niirosomonas 
h . Nitrobacter 
Family 2. Thiobacillaceae 
Tribe A. Thiobacilleae 
Genus a. Thiobacillus 



SURVEY OF CLASSIFICATIONS OF BACTERIA 21 

Order II. Metahacteriales 

Family 1. Pseudomonadaceae 
Tribe A. Spirilleae 

Genus a. Spirillum 
Tribe B. Vibrioneae 
Genus a. Vibrio 
Tribe C. Pseudomonadeae 
Genus a. Pseudomonas 
h. Azoiobacter 
Connects with Polyangiaceae and Nitrubacteriaceae 
Family 2. Bacleriaceae 
Tribe A. Aerobactereae 
Genus a. Aerobacter 
h. Escherichia 

c. Salmonella 

d. Eberthella 

e. Proteus 
Tribe B. Pasteurelleae 

Genus a. Alcaligenes 
h. Pasteurella 
Connects with Pfeifferella 

c. Hem,ophilus 
Connects with Dialisler 
Family 3. Micrococcaceae 
Tribe A. Streplococceae 

Genus a. Neisseria • 

b. Streptococcus 
Tribe B. Micrococceae 

Genus a. Micrococcus 

h. Staphylococcus 

c. Sarcina 
Connects with Algubacteria 

Subclass C. Mycobacteria 
Order I. Bacteriomycetales 
Family 1. Leptotrichaceae 
Tribe A. Acetobactereae 
Genus a. Acetobacter 
Connects with Salmonella and Tissieria 
Tribe B. Leptotricheae 
Genus a. Kurthia 

b. Lactobacillus 
Connects with Corynebacterium 

c . Leptotrichia 
Connects with Erysipelothrix 

Family 2. Bacteroidaceae 
Tribe A. Dialistereae 

Genus a. Type species Dialistereavariegata 

Connects with Distasoa 

h. Type species Dialisterea variabilis 

c. Dialisler 



22 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Connects with Hemophilus 
Tribe B. Bacieroideae 

Genus a. Type species Bacteroidea multiformis 

b. Bacter aides 
Connects with Tissieria 

c. Type species Bacteroidea fusiformis 

Order II. Bacilloviycetales 

Family 1. Bacillaceae 
Sub-family la. Aerobacilloideae 
Tribe A. Aerobacilleae 

Sub-tribe Al. Centrosporineae 
Genus a. Centrosporus 
b. Fusibacillus 
Sub-tribe A2. Aerobacillineae 
Genus a. Aerobacillus 
Tribe B. Pseudobacilleae 

Genus a. Pseudobacillus 
Sub-family lb. Bacilloideae 
Tribe A. Bacilleae 

Sub-tribe Al. Bacillineae 
Genus a. Bacillus 

h. Megatherium 
Sub-tribe A2. Astasineae 
Genus a. Astasia 
h. Plexus 
" Tribe B. Anthraceae 

Genus a. Anthrax 
Family 2. Clostridiaceae 
Sub-family 2a. Botulinoideae 
Tribe A. Botulineae 
Genus a. Botulinus 

b. Chauvoea 

c. — — Type species Botulinea saccharolytica 

d. Type species Botulinea butyrica 

Tribe B. Putrificeae 

Genus a. Putrificus 
Sub-family 2b. Clostridioideae 
Tribe A. Welchieae 

Genus a. Welchia 
Tribe B. Clostridieae 
Genus a. Clostridium 
, Order III. Actinomycetales 

Family 1. M ycobacteriaceae 

Tribe A. Actinobacilleae 

Genus a. Pfeifferella 

Connects with Pasteurella 

b. Actinobacillus 

c. Corynebacterium 

d. Erysipelothrix 
Connects with Leptotrichia 



SURVEY OF CLASSIFICATIONS OF BACTERIA 23 

Tribe B. Mycobactereae 

Genus a. Phleobacteriuni 
b. Mycobacterium 
Tribe C. Tissierieae 

Genus a. Distasoa 

b. Tissieria 

Connects with Bacteroides, Corynebacterium and Acelobacter 
Family 2. Actinomycetaceae 
Tribe A. Actinoidomycetaceae 

Genus a. Actinoidomyces 
Tribe B. Actinomycetaceae 
Genus a. Actinomyces 
Subclass D. Algobacteria 
Order I. Desmobacteriales 
Family 1. Sphaerotilaceae 

Genus a. Sphaerotilus 
Order II. Siderobacteriales 

Family 1. Chlamydotrichaceae 
Tribe A. Chlamydotricheae 
Genus a. Leptothrix 
h. Crenothrix 
Family 2. Siderocapsaceae 

Genus a. Didymohelix 
h. Siderocapsa 

c. Sideromonas 
Order HI. Thiobacteriales 

Family 1. Rhodobacteriaceae 
Sub-family la. Chromatoideae 
Tribe A. Thiocapseae 
Genus a. Thiocystis 

b. Thiosphaera 

c. Thiosphaerion 

d. Thiocapsa 

e. Thiosarcina 

f. Lamprocysiis 
Tribe B. Thiopedieae 

Genus a. Lampropedia 

h. Thioderma 
Tribe C. Amoebobacteriae 
Genus a. Amoebobacter 

h. Thiodictyon 

c. Thiothece 

d. Thiopoly coccus 
Tribe D. Chromatieae 

Genus a. Chromatium 

b. Rhabdomonas 

c. Thiospirillum 

d. Rhodocapsa 

e. Rhodcihece 



24 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Sub-family lb. Rhodobacteroideae 
Tribe A. Rhodobacteriaceae 
Genus a. Rhodobacterium 

b. Rhodobacillus 

c. Rhodovibrio 

d. Rhodospirillum 
6. Rhodosphaera 

Tribe B. Rhodocysteae 
Genus a. Rhodocystis 
b. Rhodonostoc 
Connects with Leuconosloc 
Family 2. Beggiatoaceae 
Genus a. Thiothrix 

b. Beggiatoa 

c. Thioploca 
Family 3. Achromatiaceae 

Genus a. Achrcmatium 
h. Thiophysa 

c. Thiospira 

d. Hillhousia 
Order IV. Myxobacieriales 

Family 1. Polyangiaceae 

Genus a. Chondromyces 
b. Polyangium 
Family 2. Myxococcaceae 

Genus a. Myxococcus 

Later Pribram (Klassification der Schizomyceten (Bakterien), Leipzig 
and Wien, 1933, 143 pp.) developed this classification into a suggestive out- 
line based on his experience in caring for the cultures of the Krai Collection. 
His most interesting contribution is the separation of the class of Schizo- 
mycetes into three subclasses which are based on differences in fundamental 
biological and nutritional relationships. The fourth sub-class of his earlier 
outline (the Protozoohacteria with its single order Spirochaetales) is omitted 
from this outline. The first class, Algohacteria, includes the bacteria that 
are primarily free-living in water, usually motile with polar flagellation and 
live on easily soluble foodstuffs. They are frequently surrounded by in- 
soluble secretions such as capsules, sheaths, etc., and form insoluble prod- 
ucts in their protoplasm, such as calcium, sulfur and iron compounds, and 
pigments. The class Euhacteria includes those bacteria whose normal 
habitat is the animal body or complex waste products of plant or animal 
origin. Because of adaptation to environment, these organisms are motile 
or non-motile and can utilize compounds of complex molecular structure. 
The third sub-class, Mycobacteria, is adapted to life in soil, and shows a dis- 
tinct tendency to differentiation in morphology and spore formation. 

Internationally accepted rules of nomenclature are generally followed, 
and the generic terms proposed in his earlier outline that were not formed 



SURVEY OF CLASSIFICATIONS OF BACTERIA 25 

in accordance with recommended practices are discarded. He has revived 
Ulvina Kiitzing, 1837 (status explained by Buchanan, General Systematic 
Bacteriology, 1925, p. 518) in place of Acetohacter Beijerinck and accepted 
Plocamohacteriuvi (Loewi, Wien. klin. Wchschr., 38, 1920, 730) in place of 
Lactobacillus Beijerinck, 1901. Among the spore-forming rods, he has 
accepted Bactrillum Fischer and Welchillus Heller, 1921. Malleomyces 
Hallier (Bot. Ztg., 24, 1866, 383) is used for the glanders bacillus. Anthra- 
cillus is apparentl}' new. 

The new outline has the following form : 



Class. Schizomycetes 




Subclass A. Algobacteria 


Order 1. Micrococcales 


Family 1. Micrococcaceae 


Genus a. 


Micrococcus 


b. 


Rhodococcus 


c. 


Rhodocapsa 


d. 


Thiocapsa 


e. 


Thiosphaera 


f. 


Thiosphaerion 


g- 


Thiocystis 


h. 


Lamprocysiis 


i. 


Sarcina 


J- 


Thiosarcina 


Family 2. Fediococcaceae 


Genus a. 


Pediococcus 


b. 


Lampropedia 


c. 


Thiothece 


d. 


Thiopolycoccus 


e. 


Thioderma 


f. 


Amoebomonas 


to* 


Rhodothece 


h. 


Rhodonostoc 


i. 


Thiophysa 


Order 2. Pseudomonadales 


Family 1. Pseudomonadaceae 


Genus a. 


Pseudomonas 


b. 


Rhodobacillus 


c. 


Chromatium 


d. 


Nitrosomonas 


e. 


Vibrio 


f. 


Rhodovibrio 


g- 


Myxococcus 


h. 


Spirillum 


i. 


Rhodospirillum 


J- 


Thiospira 


k. 


Thiospirillum 


Family 2. Serratiaceae 


Genus a. 


Serratia 


b. 


Hillhousia 



26 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Family 3. Nitrobacteriaceae 

Genus a. Nitrobacter 

b. Rhodobacterium 

c. Rhodocystis 

d. Didymohelix 

e. Sideromonas 

f. Siderocapsa 

g. Chondromyces 
h. Polyangium 

i. Amoebobacter 
j. Thiodictyon 
Family 4. Azotobacteriaceae 
Genus a. Rhizobium 
b. Azotobacter 
Order 3. Leptotrichales 

Family 1. Leptotrichaceae 
Genus a. Leptothrix 

b. Sphaerotilus 

c. Crenothrix 
Family 2. Clonothrichaceae 

Genus a. Clonothrix 
Order 4. Rhabdonionadales 

Family 1. Rhabdomonadaceae 
Genus a. Beggiatoa 

b. Rhabdomonas 

c. Thioploca 

d. Thiothrix 
Family 2. Spirochaetaceae 

Genus a. Spirochaeta 

b. Treponema 

c. Leptospira 

d. Cristispira 

e. Saprospira 
Subclass B. Eubacteria 

Order 1. Aerobacteriales 

Family 1. Aero6ac<enoceae 
Genus a. Aerobacter 
h. Escherichia 

c. Salmonella 

d. Eberthella 

e. Shigella 
Family 2. Pasteurellaceae 

Genus a. Pasteurella 

b. Brucella 

c. Haemophilus 

d. Neisseria 
Order 2. Plocamobacteriales 

Family 1. Streptococcaceae 
Genus a. Streptococcus 

Family 2. Ulvinaceae 
Genus a. Proteus 
h. Kurthia 



SURVEY OF CLASSIFICATIONS OF BACTERIA 27 

c. Ulvina 

d. Plocarnobacterium 

e. Leptolrichia 
Family 3. Bacteroidaceae 

Genus a. Dialister 

b. Aerobacteroides 

c. Bacteroides 

d. Fusobaderium 
Subclass C. Mycobacteria 

Order 1. Bacillales 

Family 1. Bacillaceae 
Genus a. Bactrillum 
h. Aerobacillus 

c. Bacillus 

d. Anthracillvis 
Family 2. Closlridiaceae 

Genus a. Clostridium 
b. Welchillus 
Order 2. Mycobacteriales 

Family 1. Mycobacteriaceae 
Genus a. Malleomyces 



b. 


Actinobacillus 


c. 


Corynebacterium 


d. 


Erysipelothrix 


e. 


Mycobacterium 


f. 


Distasoa 


g- 


Tissieria 


mily 2. . 


4 ctino mycetaceae 


Genus a. 


Actinomycoides 


b. 


Actinomyces 



Janke (Cent. f. Bakt., II Abt., 80, 1930, 481) reprints the earlier outline 
prepared by Pribram (1929) and, after commenting on Lehmann and 
Neumann's (1927) outline, proposes an outline which is slightly modified 
from his own previous (1929) outline. Two new subgeneric terms are used, 
Anaerobacillus and Euhacterium. The sub-genus Aerobacillus is apparently 
not the same as Aerohacillus Donker (Inaug. Diss., Delft, 1926), nor as 
Aerobacillus Pribram (Jour. Bact., 18, 1929, 361). 

Family T. Micrococcaceae 
Genus 1. Micrococcus 





Divided into 2 sections. 


2. 


Neisseria 


3. 


Streptococcus 




Divided into 4 sections. 


4. 


Sarcina 




Divided into 2 sections. 


mily II. 


Bacillaceae 


Genus 1. 


Bacillus 


Sub-genus a. Anaerobacillus or 1 



Divided into 6 sections. 



28 MANUAL OF DETERMINATIVE BACTERIOLOGY 





b. Aerobacillus or better Eubacillus 




Divided into 10 sections. 


Family III. 


Bacteriaceae 


Genus 1. 


Bacterium 


Sub-genus a. Pseudomonas 




Divided into 6 sections. 




b. Eubacterium 




Divided into 11 sections. 




c. Trichobacterium 




Divided into 6 sections. 


Genus 2. 


Fusiformis 


Family IV. 


Corynobacteriaceae 


Genus 1. 


Mycobacterium 


2. 


Pfeifferella 


3. 


Erysipelothrix 


4. 


Corynobacterium 


5. 


Actinomyces 


Family V. 


Spirillaceae 


Genus 1. 


Microspira or Vibrio 


Sub-genus a. Microspira 




b. Spirosoma 


Genus 2. 


Spirillum 


Sub-genus a. Spirella 




h. Dicrospirillum 


Family VI. 


Spirochaetaceae 


Genus 1. 


Spirochaeta 


2. 


Cacospira 


3. 


Entomospira 


4. 


Treponema 


5. 


Cristispira 


6. 


Saprospira 


7. 


Leptospira 


Family VII. 


Desmobacteriaceae 




As in 1929 outline. 


Family VII] 


'.. Myxobacteriaceae 




As in 1929 outline. 



Kluyver and Van Niel (Cent. f. Bakt., II Abt., 94, 1936, 369) have 
developed an outline classification in which they indicate four lines of 
development from the simplest form of cell that is existent and conceivable, 
the sphere. They assign family rank to each of these four groups of bac- 
teria, placing the lophotrichous (and related non-motile) rod-shaped bac- 
teria first (Pseudomonadaceae) . This is followed by the family of spherical 
bacteria (Micrococcaceae) and the family of permanently non-motile, rod- 
shaped bacteria (Mycohacteriaceae) . The final family includes the peritri- 
chous (and related non-motile) rod-shaped bacteria, the Bacteriaceae. 
These are grouped in the tribes of each family in accordance with their 
fundamental metabolism as photo-autotrophic,photo-heterotrophic,chemo- 
autotrophic and chemo-heterotrophic. Their outline follows: 



SURVEY OF CLASSIFICATIONS OF BACTERIA 29 

Family A. Pseudomonadaceae 
I. Tribe Spirilleae 



Genus 1. 


Thiospirillum 


2. 


Phaeospirillum 


3. 


Rhodos'pirillnvi 


4. 


Sulfospirillum 


5. 


Spirillutn 


II. Tribe Vibrioneae 


Genus 1. 


Chromaiium 


2. 


Rhodovibrio 


3. 


Didymohelix 


4. 


Vibrio 


5. 


Desulfovibrio 


III. Tribe Pse 


udomonadeae 


Genus 1. 


Thiothece 


2. 


Phaeomonas 


3. 


Rhodomonas 


4. 


Sulfomonas 


5. 


Sideromonas 


6. 


Nitroso7nona!< 


7. 


Nitrobader 


8. 


Acetobader 


9. 


Pseudotnonas 


10. 


Rhizobium 


11. 


Azotobader 


12. 


Listerella 


13. 


Aeromonas 


14. 


Zymomonas 


15. 


Methanobaderium 


Family B. Micrococcaceae 


IV. Tribe Mi 


crucocceae 


Genus 1. 


Chlorobium 


2. 


Thiopoly coccus 


3. 


Rhodococcus 


4. 


A chromaiium 


5. 


Siderocapsa 


6. 


Nitrosococc7is 


7, 


Neisseria 


8. 


Micrococcus 


9. 


Veillonella 


10. 


Peptococcus 


11. 


Methanococcus 


V. Tribe Sarcineae 


Genus 1. 


Thiopedia 


2. 


Thiosarcina 


3. 


Gaffkya 


4. 


Sarcina 


5. 


Zymosarcina 


6. 


Butyrisarcina 


7. 


Methanosarcina 



30 MANUAL OF DETERMINATIVE BACTERIOLOGY 

VI. Tribe Sporosarcineae 
Genus 1. Sporosarcina 
VII. Tribe Sireplococceae 

Genus 1. Peptostreplococcus 

2. Streptococcus 

3. Betacoccus 
Family C. M ijcobacteriaceae 

VIII. Tribe Corynebacterieae 

Genus 1. Corynebacterium 





2. Fusiformis 




3. Propionibacterium 




4. Streptobacterium 




5. Betabacterium 


IX. 


Tribe Mycobacterieae 




Genus 1. Mycobacterium 




2. Thermobacterium 


fiily 


D. Bacteriaceae 


X. 


Tribe Bacterieae 




Genus 1. Kurihia 




2. Alcaligenes 




3. Bacterium 




4. Aerobacter 


XI. 


Tribe Bacilleae 




Genus 1. Bacillus 




2. Aerobacillus 




3. Zymobacillus 




4. Clostridium 




5. Peptoclostridium 



Some old names are displaced by new descriptive terms : Phaeospirillum 
Sulfospirilluin, Desulfovibi'io, Phaeomonas, Aeromonas, Zymomonas, Me- 
thanobacterium, Methanococcus, Meihanosarcina, Butyrisarcina, Peptococcus, 
Peptostreplococcus, Zymobacillus. Rhodomonas is not used in the same sense 
as Rhodomonas Orla-Jensen (Cent. f. Bakt., II Abt., £2, 1909, 331 and 334), 
the latter being a synonym of Chromatium Perty (Zur Kenntniss kleinster 
Lebensformen, 1852). Sulfomonas is indicated as new and as a synonym 
of Thiobacillus Beijerinck (Cent. f. Bakt., II Abt., 11, 1904, 598) although 
the same term is used by Orla-Jensen (loc. cit.). Three new terms are 
accepted : C/i/oro6zMm Nadson (Bull. Jard. Bot. St. Petersburg, 6, 1906, 184), 
Zymosarcina Smit (Die Garungssarcinen, Jena, 1930) and Peptoclostridium 
CDonker, Inaug. Diss., Delft, 1926). 

Rahn (Cent. f. Bakt., II Abt., 96, 1937, 273) has reviewed the characters 
of the species of Eubacteriales included in the fourth edition of this Manual. 
He places 146 of the spore-forming species in a Sub-order A. Endosporales 
with a single family, and 536 of the species of non-spore-forming rods in a 
Sub-order B. Asporales in seven families. Unclassifiable species (total 224) 
are placed in a temporary eighth family Bacteriaceae. His outline follows: 



SURVEY OF CLASSIFICATIONS OF BACTEEIA 31 

Order Eubacteriales 

Suborder A. Endosporales 
Family I. Endosporaceae 
Genus 1. Bacillus 

2. Aerobacillus 

3. Clostridium 
Suborder B. Asporales 

Family I. Gramoxidaceae 

Genus 1. Micrococcus {inclndmg Staphylococcus, Gaff- 
kya, Rhodococcus and most of the species 
of Sarcina) 
2. Kurthia 
Family II. Gramanoxidaceae 
Tribe a. Streptococceae 

Genus 1. Streptococcus (including Diplococcus) 

2. Leuconostoc 

3. Peptostreptococcus 
Tribe b. Lactobacilleae 

Genus 4. Lactobacillus (including part of Bacter- 
oides) 
5. Propionibacterium 
Tribe c. Sarcineae 

Genus 6. Zymosarcina 

7. Butyrisarcina 

8. Methanosarcina 
Family III. Neissereaceae 

Genus 1. Neisseria 
2. Veillonella 
Family IV. Protobacteriaceae 
Tribe a. Protobacterieae 

Genus 1. Carboxydomonas 
2. Methanomonas 
Tribe b. Nitrobacterieae 
Genus 1. Nitrosomonas 

2. Nitrobacter 

3. Nitrosococcus 
Family V. Enterobacteriaceae 

Genus 1. Enter obacter (including Escherichia, Sal- 
monella, Aerobacter, Klebsiella, Proteus, 
Erwinia, Eberthella, Shigella, and parts 
of Serratia, Pseudomonas, Flavobacterium 
and Achromobacter) 
Family VI. Pseudomonadaceae 

Genus 1. Pseudomonas (includes Phytomonas and 
other lophotrichous types only) 

2. Vibrio 

3. Spirillum 

4. Acetobacter 

5. Azotobacter 

6. Rhizobium 



32 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Family VII. Parvobacieriaceae 
Genus 1. Brucella 

2. Pasteur ella 

3. Hemophilus (including Dialister) 
Family VIII. Bacteriaceae 

Unclassifiable genera including Alcaligenes 
and Protaminobacier ; some species from 
each of the following genera, Achromobac- 
ter, Chromobacterium, Cellulomonas, Bacter- 
oides, Flavobacterium, Phytomonas, Pseudo- 
monas, Serratia; and three species from the 
Family Nitrobacteriaceae. 

One of the generic terms used in this outline is new, i.e., Enterohacter. 
Two other generic terms, Fluorescens and Erythrohacterium, are proposed 
incidentally (p. 284). The first includes the peritrichous forms included in 
the Manual under Pseudomonas and the second includes those red, non- 
spore-forming rods that are not included in Serratia. In another footnote 
(p. 281) a substitute, Virgula, is suggested for Enterohacter. Emphasis is 
placed on sporulation, Gram stain, and oxygen demand as the most im- 
portant characters aside from cell form and flagellation. 

Prevot, as an outgrowth of his studies on anaerobes with Weinberg 
(Weinberg, Nativelle and Prevot, Les microbes anaerobies, 1937, 1186 pp., 
Paris), has written a series of papers in which he has developed a classifica- 
tion of anaerobic bacteria (Ann. Sci. Nat., 10 Ser., 15, 1933, 23-260; Ann. 
Inst. Past., 60, 1938, 285-307; 61, 1938, 72-91; 64, 1940, 117-125). The 
conclusions reached in these studies are summarized in his Manual de Clas- 
sification et de Determination des Bacteries Anaerobies, Monographic de 
ITnstitut Pasteur, Paris, 1940, 223 pp. He regards the bacteria as com- 
prising a kingdom, Schizomyceles, intermediate bfetween the animal and 
plant kingdoms and notes the presence of strict anaerobes in at least three 
of the seven orders recognized in the 5th edition of the Manual. These 
orders he regards as classes. The genus Bacteroides Castellani and Chalm- 
ers (Manual of Trop. Med., 3rd ed., 1919, 959) type species, Bacteroides 
fragilis, is dropped (Ann. Inst. Past., 60, 1938, 288), and several new terms 
are proposed for the organisms included by Castellani and Chalmers and 
later investigators in the genus. Among the new generic names is Ristella 
which is based on Ristella fragilis, the species used by Castellani and 
Chalmers as the type species for Bacteroides. 

The complete outline classification developed bj^ Prevot in his Mono- 
graph (loc. cit., p. 17) is given below: 

Kingdom. Schizomyceles Nageli 
Class I. Eubacteriales 

Sub-Class I. Non sporogenous Eubacteriales 
Order I. Micrococcales 



SUKVET OF CLASSIFICATIONS OF BACTERIA 33 

Family I. Neisseriaceae 
Tribe I. Neisserieae 

Genus a. Neisseria 
Tribe 2. Veillonelleae 
Genus a. Veillonella 
Family 2. Micrococcaceae 
Tribe 1. Streplococceae 
Genus a. Diplococcus 
b. Streptococcus 
Tribe 2. Staphylococceae 
Genus a. Gaffkia 

b. Staphylococcus 
Tribe 3. Micrococceae 
Genus a. Sarcina 

b. Micrococcus 
Order II. Bacteriales 
Family 1. Ristellaceae 
Genus a. Ristella 

b. Pasteur ella 

c. Dialister 

d. Zuberella 

e. Capsularis 
Family 2. Bacteriaceae 

Genus a. Eubacterium 

b. Catenabacterium 

c. Ramibacterium 

d. Cillobacterium 
Order III. Spirillales 

Family 1. Vibrionaceae 
Genus a. Vibrio 
Sub-class II. Sporogenous Eubacteriales 
Order I. Clostridiales 

Family 1. Endosporaceae 

Genus a. Endosporus 
b. Paraplectrum 
Family 2. Clostridiaceae 
Genus a. Infiabilis 

b. Welchia 

c. Clostridium 
Order II. Plectridiales 

Family 1. Terminosporaceae 

Genus a. Terminosporus 
b. Caduceus 
Family 2. Plectridiaceae 

Genus a. Pleclridium 
b. Acuformis 
Order III. Sparovibrionales 
Family 1. Sporovibrionaceae 
Genus a. Sporovibrio 



34 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Class II. Adinomycetales 

Family 1. Spherophoraceae 

Genus a. Spherophorus 





b. 


Spherocillus 




c. 


Fusiformis 




d. 


Fusocillus 




e. 


Leptotrichia 


Family 2. 


Actinomycetaceae 


Genus a. 


Actinobacterium 




b. 


Bifidibacterium 




c. 


Corynebacterium 


Class III. Spirochetales 




Family 1. 


Spirochaetaceae 


Genus a. 


Treponema 




b. 


Borrelia 



In this outline, there are minor modifications in the names and in endings 
given to the orders and tribes as compared with those given in his prehm- 
inary papers. In the Order Micrococcales, Leuconostoc has been dropped 
as a genus of the tribe Strepiococceae and Rhodococcns has been dropped as 
a genus of the Tribe Staphylococceae. Veillonella proposed by Pr^vot as a 
new genus in 1933 {loc. cit., p. 70) is included as a genus in the Family 
Neisseriaceae. The spelling of Gajjkya is changed to Gaffkia. In the first 
of Prevot's papers published in 1938 {loc. cit.), he proposes the following 
new genera in the Order Bacteriales: Ristella, Zuherella, Capsularis, Euhac- 
terium, Catenabacteriuni, Ramihacterivm and Cillohacterium. In the same 
paper he also proposes the following new genera in the Order Actinomy- 
cetales: Spherophorus, Spherocillus, Fusocillus, Pseudoleptothrix (withdrawn 
in 1940 in favor of Leptotrichia Trevisan). He also accepts one genus 
Actinohacterium (Haas, Cent. f. Bakt., I Abt., Orig., ^0, 1906, 180) not 
previously mentioned in this discussion. With the single change noted 
{Pseudoleptothrix to Lepototrichia) , the outlines of the genera in the orders 
Bacteriales and Actinomycetales remains in the 1940 outline as it was given 
in 1938. 

In the outline given in Prevot's Monograph {loc. cit., p. 17) one change is 
made in the generic terms recognized in the r)rder Clostridiales from those 
recognized in his second paper published in 1938, The genus name 
Palmula proposed in 1938, having been found to l3e invalid because of prior 
use for a genus of Protozoa, is changed to Acuformis. Other generic names 
which appeared for the first time in the 1938 outline are Endosporus, 
Inflahilis, Terminosporus and Caduceus. ]Velchia proposed by Pr^vot in 
^933 {loc. cit., p. 44) was previously proposed by Pribram (Jour. Bact., 18, 
1929, 374) for the same group of anaerobic spore-forming rods. A third 
order, Sporovibrionales, is proposed by Prevot in his Monograph {loc. cit., 
p. 15) to include the family Sporovibrionaceae (Ann. Inst. Past., 64, 1940, 



SURVEY OF CLASSIFICATIONS OF BACTERIA 35 

119). This order and family include a single genus Sporovihrio Starkey 
(Arch. f. Microb., 9, 1938, 300) syn. Desulfovibrio Kluyver and Van Niel 
(Cent. f. Bakt., II Abt., 94, 1936, 389). Two genera {Treponema and 
Borrelia) of Spirochaetales are listed by Prevot in his Monograph {loc.cit., 
p. 16) as including anaerobic species. 

Stanier and Van Niel (Jour. Bact., 4^, 1941, 437-466) have proposed a 
rearrangement of the classification outline as indicated below: 

Kingdom Monera 

Division I Myxophyta (Blue-green algae) 

Division II Schizomycetae (Bacteria) 
Class I Eubacteriae 

Order I Rhodobacteriales 

Order II Eubacteriales 

Order III Adinowycetales 

Class II Myxobacteriae 

Order I Myxobacteriales 

Class III Spirochaetae 

Order I Spirochaetales 

Ajjpendix to Division Schizomycetae 

Group I Includes two families, Leptotrichaceae and Crenothri- 

caceae 
Group II Achromaiiaceae 

Group III Pasleuriaceae (Includes three genera, Pas(eicria,Hypho- 
viicrobium and Blastocaulis) 

The genera Mycobacterium, Corynehacterhim, Erysipelothrix, Leptotrichia, 
Nevskia, GallioneUa, Caulohacter, Thiospira, Siderocapsa and Sideromonas 
are placed in Eubacteriales. Two genera not previously discussed in this 
review whose relationships to other bacteria have recently been clarified 
are Sporocytophaga Stanier (Jour. Bact., 40, 1940, 629) and CTjtophaga 
Winogradsky (Ann. Inst. Past., 43, 1929, 578). 

This rearrangement has been carried out by including the organisms 
placed in the Order Caulobacteriales Henrici and Johnson (Jour. Bact., 30, 
1935, 61-93) in the Order Eubacteriales (Buchanan, Jour. Bact., 3, 1917, 
162). The genera of the Order Chlamydobactcriales Buchanan {loc. cii.) 
are transferred to an appendix or are dropped {Clonothrix) as belonging to 
the blue-green algae. Three of the remaining five orders are raised to the 
rank of classes, one of which (Eubacteriae) includes three orders Rhodo- 
bacteriales (Pringsheim, Lotos, 71, 1923, 351), Eubacteriales (Buchanan, 
loc. cit.) and Actinomijcetales (Buchanan, loc. cit.). Rhodobacteriales in- 
cludes the sulfur purple, the non-sulfur purple and the green bacteria, the 
colorless sulfur bacteria {Beggiatoaceae) being transferred to the Myxophyta 
with the change of the name of the Order from Thiobacteriales Buchanan 
(loc. cit.) to Rhodobacteriales Pringsheim {loc. cit.). 



36 MANUAL OF DETERMINATIVE BACTERIOLOGY 

The outline classification below is proposed by the Editorial Board of the 
Manual for use in the present (6th) edition of the Manual. It is based on 
those developed by Bergey et al. in earlier editions. These, in turn, were 
based on the outline classifications developed bj^ Buchanan (Jour. Bact., 1, 
1916, 591 ; 2, 1917, 155 ff.; 5, 1918, 27 ff.) and Winslow et al. (Jour. Bact., 5, 
1920, 191). 

Phylum Schizo'phyta 

Class I. Schizophyceae 
Class II. Schizomycetes 
Order I. Eubacteriales 

Sub-Order I. Eubacteriineae (includes Corynebacteriaceae) 
Sub-Order II. Caulobacteriineae 
Sub-Order III. Rhodobacteriineae 
Order II. Actinomyceiales (includes Mycobacterium, Actinomyces, 

and related genera) 
Order III. Chlamydobacteriales 
Family I. Leptotrichaceae 
Family II. Crenothrichaceae 
Family III. Beggiatoaceae 

Appendix Achromatiaceae 
Order IV. Myxobacteriales 
Order V. Spirochaetales 
Supplement: Groups whose relationships are uncertain. 

Group I. Order Rickettsiales . Group II. Order Virales. Group III. 
Family Borrelomycetaceae. 

In this, the arrangement of Schizomycetes as a class coordinate with 
Schizophyceae, both belonging to a phylum Schizophyta of the plant king- 
dom, is maintained as before. The number of orders is reduced from seven 
as given in the fifth edition of the Manual to five, through recognition of 
the fact that the rigid, unicellular, sometimes branching but never truly 
mycelial nor filamentous organisms belonging to three of the previously 
recognized orders are presumably more closely related to each other than 
they are to the organisms in the four remaining orders. The family Coryne- 
bacteriaceae has been transferred from the order Actinomyceiales to Eubac- 
teriales. 

The colorless, filamentous, sulfur bacteria (Beggiatoaceae) have been 
placed in the order Chlamydobacteriales with the other filamentous bacteria 
that are clearly related to the blue-green algae. While this marks the 
greatest deviation from the outline previously used, and separates these 
colorless sulfur bacteria from the purple sulfur bacteria placed in Rhodo- 
bacteriineae, it is in accordance with the arrangement accepted by Lehmann 
and Neumann (Bakt. Diag., 4 Aufl., 2, 1907, 598), Pringsheim (Lotos, 71, 
1923, 307) and others. Rhodobacteriineae is also limited to the purple and 
green bacteria as suggested by Pringsheim Qoc. cit.) and accepted by Kluy- 
ver and Van Niel (loc. cit.), by Stanier and Van Niel (loc. cit.) and others. 



SURVEY OF CLASSIFICATIONS OF BACTERIA 37 

The Rickettsiales and Borrelomycetaceae are placed in a supplement as 
their relationships are still obscure. Several authors would place them 
near some of the organisms now placed in Pasteurella and Haemophilus. 
The viruses {Virales) whose nature and relationships are still more ob- 
scure are also placed in a supplemental group. 

Although this outline maintains the simplicitj' that distinguished its 
predecessors, and provides places for all tj^pes of microorganisms thus far 
described that may properly be grouped under the fission fungi, it should 
not be regarded in any sense as final. An attempt has been made to express 
natural relationships, but these are so frequently obscure or unknown that 
in many places utilitarian considerations have prevailed. In some places, 
groups of known doubtful significance have been allowed to stand as they 
are out of a desire not to make unnecessar}^ changes. It has appeared 
desirable to be conservative in making changes in the outline as used 
previously. 

Addenda: After the above was in page proof, it was discovered that 
reference to the outline classification of Gieszczykiewicz (Bull. Adad. 
Polonaise d. Sci. et d. Lettres, CI. Sci. Math, et Nat., S^r. B., 1939, 27 pp.) 
had inadvertently been omitted. This outline has some features like the 
outline that Lehmann and Neumann used in 1927 (see p. 17) and some 
like the outline used in the 4th ed. of the Manual. 

The genus Bacterium is retained as in the Lehmann and Neumann out- 
line for Gram-negative, non-spore-forming, peritrichous or polar flagellate 
rods. Twelve sub-genera are recognized and these bear subgeneric scien- 
tific names that are much the same as those used for genera in the 4th ed. 
of the Manual. A new subgeneric name Enterohaderium (see Entero- 
hader Rahn) is proposed to cover the genera Escherichia, Aerohader, 
Klebsiella, Salmonella, Eherthella and Shigella. Loefflerella previously used 
by Gay et al. (Agents of Disease and Heat Resistance, Indianapolis, 1935, 
782) is here also used as a subgeneric name for the glanders bacillus; and 
Chromobaderium is used for the organisms more properly placed in Serratia 
Bizio. 

Corynebaderium is transferred from, the order Aciinomycetales to Eubac- 
teriales and the famih^ Corynebaderiaceae is made to include Lactobacillus, 
Erysipelothrix and Fusobadcrium. .Among the Spirochadales, the genus 
name Spirochaeta is displaced by a new generic term, Ehrenbergia, and is 
itself used to displace Borrelia. 

A seventh order RicJcdtsiales is proposed to include two families: Rickeit- 
siaceae with one genus Rickettsia da Rocha Luna (Berl. klin. Wchnschr., 
1916, 567); and Bartonellaceae with the genera, Bartonella Strong, Tyzzer, 
Bnies, Sellards and Gastiabuni (Jour. Amer. I\Ied. Assoc, 61, 1913, 
1713), and Grahamella Brumpt (Bull. Soc. Path. Exot., 4, 1911, 514). 



38 MANUAL OF DETERMINATIVE BACTERIOLGY 

During 1945, Soriano (Ciencia e Investigacion, 1, 1945, 92-94 and 146- 
147; Rev. Argentina de Agronomia, 12, 1945, 120) proposed an arrange- 
ment of the Class Schizomycetes in which he recognizes a new Order, Flexi- 
hacteriales, to inckide the [families Cyiophagaceae and Beggiatoaceae and 
an entirely new Family Flexibacteriaceae containing a single genus Flexi- 
hader. The latter includes five newly recognized species of flexuous bacteria 
as follows: Flexihacter flexilis, type species, F. elegmis, F. giganteus, F . al- 
huminosus and F . aureus. 

The outline given below shows how this new order and new family are 
fitted by Soriano into the classification used in the fifth edition of the 
Manual. 

Class Schizomycetes 
Subclass Eubacteria. Rigid cells. Subclass Flexibacteria. Flexuous cells. 
Order I. Eubacteriales Order VI. Flexihacter iales 

Order II. Caulobacferialei< Family I. Cytophagaceae 

Order III. Rhodobacter iales Family II. Beggiatoaceae 

Order IV. Actinomycetales Family III. Flexibacteriaceae 

Order V. Chlamydobacteriales Order VII. M yxobacteriales 

Order VIII. Spirochaetales 

Prevot (Ann. Inst. Past. 72, 1946, 1) has developed his classification of 
Class Actinomycetales, subdividing it into orders and including several 
genera not recognized in his 1940 outline. This classification is as follows: 

Class Actinomycetales 

Order I. Actinubacteriales. New order. Not acid-fast 
Family I. Spherophoraceae. Family II. Actinomycetaceae. 

Genus I. Spherophorus Genus I. Actinomyces 

Genus II. Harerhillia Genus II. Proactinomyces 

Genus III. Spherocillus Genus III. Corynchacterium 

Genus IV. Fusiformis Genus IV. Actinobacterium 

Genus V. Fusocillus Genus V. Bifidibacterium 

Genus VI. Leptotrichia Genus VI. Erysipelothrix 

Order II. Mycobacter iales. New order. Acid-fast 
Genus I. Mycobacterium 

This classification differs from that used m this edition of the Manual 
in that it places several genera of Gram-negative organisms in Actinomy- 
cetales. These are Spherophorus, Havcrhillia, Spherocillus, Fusiformis and 
Fusocillus, all of which are included here under Parvohacteriaceae. Lep- 
totrichia which Prevot regards as Gram-negative is generally accepted as 
being a Gram-positive group. It is discussed in this edition of the Manual 
in connection with the genus Lactobacillus. 



HOW BACTERIA ARE NA:VIED AND IDENTIFIED* 

Some principles of taxonomy and nomenclature. "Taxonomy is that 
branch of biology that deals with the orderly arrangement of plants and 
animals" (Johnson, Taxonomy of the Flowering Plants, New York, 
1931, p. 3). 

The necessity for applying names to species or kinds of bacteria is self- 
evident. It is highly desirable that the name applied to an organism by 
one person should be understood b}^ others. It is further desirable that as 
far as practicable all individuals use the same name for the same kind of 
organism. It is helpful, therefore, if there can be an agreement regarding 
the method of naming organisms, and as to the correct name for each 
organism. The term nomenclature is applied to the naming of plants and 
animals, and under this term may be included all discussions as to methods 
of naming and correctness of particular names. 

It is not enough that bacteria be named. Some method of classification 
of the bacteria is essential if the names are to be rendered accessible and 
available, and identification of unknown forms be made possible. Taxo- 
nomy is that Ijranch of biology which treats of classification in accordance 
with a convention or law. It is apparent that taxonomy must be depend- 
ent in part for its satisfactory development upon nomenclature. Even 
though there may not be agreement among bacteriologists as to the exact 
classification that is to be used, nevertheless it is highly desirable that there 
be agreement as to some of the fundamental characteristics of satisfactory 
biological classifications in general. 

What kinds of names are used. Two kinds of names are commonly 
given to the different kinds of plants and animals, the common, provincial, 
vernacular or casual names on the one hand and the international or scien- 
tific names on the other. These should be carefully differentiated , and their 
respective advantages and disadvantages noted. 

It is inevitable, and on the whole probably' desirable, that for each kind 
of familiar animal or plant in each language there will be coined a name. 
Usually the name for the same organism will be different in each language. 
For example, we have in English Oak, in German Eiche, in Latin Qucrcus, 
etc. For many uncommon kinds, however, there may be no such vernacu- 
lar names developed. There have been, of course, many casual or vernac- 
ular names given to kinds of bacteria. In English we speak of the tubercle 
bacillus, the typhoid germ, the gonococcus, the Welch bacillus, the golden 

* Contributed by Prof. R. E. Buchanan, Iowa State College, Ames, Iowa, January, 
1934; revised, March, 1943. 

39 



40' MANUAL OF DETERMINATIVE BACTERIOLOGY 

pus COCCUS, and many others. Similarly, we find in German Typhus- 
bazillen and in French bacille typhique, enterococcus, etc. The use of 
these common names offers certain advantages. It does away frequently 
with the necessity of repeating longer and more formal scientific names. 
Not infrequently scientific names may be adopted into a language, and 
converted into vernacular names. For example, the English name aster 
and the scientific generic name Aster are applied to the same group. This 
is frequently a convenience, but there are also some difficulties, which will 
be emphasized below. 

In contrast to common, vernacular or casual names, the scientific name 
for each kind of organism (each plant or animal) is supposed to be the same 
in all countries and in all languages. When such a scientific name is used, 
no question should arise in any language as to what organism is intended. 
The names thus applied are supposed to conform to certain general rules 
that have been formulated by international agreement. Obviously the use 
of such names is advantageous whenever one is desirous of accuracy, and of 
being definitely understood in all languages. It is further evident that in 
all questions relating to taxonomy and classification it is highly desirable 
that the scientific names be used. 

International rules for nomenclature. In order that there be an inter- 
national set of scientific names, it is essential that there be an international 
agreement as to the rules which should govern their creation. Both of the 
great groups of biologists, the botanists and the zoologists, have met in 
numerous international congresses in which delegates were accredited by 
the great botanical and zoological societies, museums, and educational 
institutions of the world. Codes of nomenclature designed to tell how names 
shall be manufactured and used, and how to tell which of two or more names 
that have been used is correct, have been developed by each of these groups. 
These codes or lists of rules and recommendations are quite similar in 
essentials for botany and zoology, although they differ in some details. 

The question arises: Are either or both of these codes satisfactory or 
adaptable to the use of bacteriologists. Three views have been expressed 
by various writers. Some few have suggested that the naming of bacteria 
cannot well conform to the approved international rules as their classifica- 
tion involves considerations not familiar to botanists and zoologists gen- 
erally. The second group, also a very small one, has insisted that uni- 
cellular forms of life are neither plants or animals, but protista, and that 
taxonomic rules, etc., should be distinct for this group and coordinate with 
the corresponding rules for plants and for animals. 

The third view, more commonly expressed, is that the bacteria are 
sufficiently closely related to the plants and animals, so that (in so far as 



HOW BACTERIA ARE NAMED AND IDENTIFIED 4i 

they apply) the mternational agreements of the botanists (or zoologists) 
should be used as a basis for naming them. 

International opinion on this topic was finally crystallized by resolutions 
adopted by the First International Congress of the International Society 
for Microbiology held in Paris in 1930 and by the Fifth International 
Botanical Congress held in Cambridge, England in the same year. 

The resolutions unanimously adopted by the plenary session of the 
International Society for ^Microbiology were in part as follows : 

"It is clearly recognized that the living forms with which the micro- 
biologists concern themselves are in part plants, in part animals, and in part 
primitive. It is further recognized that in so far as they may be applicable 
and appropriate the nomenclatural codes agreed upon by International 
Congresses of Botany and Zoology should be followed in the naming of 
micro-organisms. Bearing in mind, however, the peculiarly independent 
course of development that bacteriology has taken in the past fifty years, 
and the elaboration of special descriptive criteria which bacteriologists 
have of necessit}^ developed, it is the opinion of the International Society 
for Microbiology that the bacteria constitute a group for which special 
arrangements are necessary. Therefore the International Society for 
Microbiology has decided to consider the subject of bacterial nomenclature 
as a part of its permanent program." 

The International Societj^ of ]Microbiologists established a permanent 
Nomenclature Committee to pass upon suggestions and to make recom- 
mendations. This committee is composed of members from all participat- 
ing nations. Two secretaries were named, one (Dr. St. John-Brooks of the 
Lister Institute, London, England) to represent primarily medical and 
veterinary bacteriology, and one (Dr. R. S. Breed, New York State Agri- 
cultural Experiment Station, Geneva, New York, U. S. A.) to represent 
other phases of bacteriology. 

The cooperation of the International Botanical Congress was solicited 
in the naming of this committee. The resolutions were approved by the 
Section on Bacteriology of the Botanical Congress and the Congress itself 
incorporated into the Botanical Code certain special provisions relating to 
the bacteria. It also specificallj^ recognized the International Committee 
as the body to prepare recommendations relating to bacterial nomenclature. 

It is apparent, therefore, that there has been international agreement 
(in so far as this can be achieved) that bacteriologists should follow the 
botanical or zoological codes in the naming of bacteria to the extent they 
are applicable, and that exceptions or new problems should be presented 
to the International Committee. 

These rules are so important in determining the validity of bacterial 
names that the rules of the Botanical Code are included in somewhat 



42 MANUAL OF DETERMINATIVE BACTERIOLOGY 

abridged form in the section that follows this introduction. Any student 
who has occasion to name a new species or a new genus or determine the 
validity of a name should familiarize himself with these rules and recom- 
mendations. 

An effort has been made in the present volume to use nomenclature in 
conformity with these rules. 

Some general principles of nomenclature. Every student of bacteriology 
should be familiar with certain rules of nomenclature if he is to use names 
intelligently. If he wishes to correct names improperly used or if he desires 
to name new species, there are additional rules which he must observe. 

1. Each distinct kind of bacterium is called a species. 

2. To each distinct species a name is given consisting usually of two 
Latin words, as Bacillus suhtilis. 

3. The first word is the name of the genus or group to which the organism 
belongs. It is always written with a capital letter. It is a Latin or Greek 
word, or a new word compounded from Latin or Greek roots, or it may be 
derived from some other language ; but this is important, whatever its origin 
when used as a generic name it must be regarded and treated as a Latin 
noun. If it is a word not found in classic Latin, it is regarded as modern 
Latin. Some generic names in bacteriology which are Latin or formed 
from Latin roots are Bacillus (masculine) a small rod; Cristispira (fem- 
inine) a crested spiral ; Lactobacillus (masculine) a milk small rod ; Sarcina 
(feminine) a packet or bundle. Many others are words from the Greek or 
compounded from Greek roots, with the words transliterated into Latin 
letters and endings in conformity with Latin usage ; words of Greek origin 
are Micrococcus (masculine) a small grain (sphere); Bacterium (neuter) a 
small rod; Clostridium (neuter) a small spindle; Corynehacterium (neuter) 
clubbed small rod; Actinomyces (masculine) ray fungus. Other generic 
names have been given in honor of persons or places as Bcggiatoa (feminine), 
Borrelia (feminine), Eberthella (feminine), Pasteurella (feminine), Erwinia 
(feminine), Zopfius (masculine). 

4. The second word in the scientific name is a specific epithet. It is not 
capitalized except that certain authors capitalize species names derived 
from proper nouns. 

It may be : 

(a) An adjective modifying the noun, and indicating by its ending agree- 
ment with the generic name in gender, as Bacterium album (white Bac- 
terium), Bacillus alb us (white Bacillus), Sarcina alba (white Sarcina), 
Eberthella dispar (the different Eberthella) , Bacterium variabile (the variable 
Bacterium), Brucella melitensis (the maltese Brucella), Bacillus teres (the 
rounded Bacillus), Bacillus graveolens (sweet- smelling Bacillus). 



HOW BACTERIA ARE NAMED AND IDENTIFIED 43 





Typical adjectives 




Masculine 


Feminine 


Neuter 


albus 


alba 


album 


niger 


nigra 


nigrum 


tener 


tenera 


tenerum 


acer 


acris 


acre 


variabilis 


variabilis 


variabile 


dispar 


dispar 


dispar 


coccoides 


coccoides 


coccoides 


aerogenes 


aerogenes 


aerogenes 



(b) An adjective in the form of the present participle of a verb, as 
Clostridium dissolvens (the dissolving Clostridium, in the sense of the 
Clostridium which is able to dissolve), Bacillus adhaerens (the adhering 
Bacillus), Acetobacter ascendens (the climbing Acetobacter) , Bacillus esterifi- 
cans (the ester-producing Bacillus). The endings for present participles 
used as adjectives are the same for all genders. The past participle is used 
occasionally, as in Pseudomonas aptata (the adapted Pseudotnonas), Spiril- 
lum attenuatum (the attenuated Spirillum). 

(c) A noun in the genitive (possessive) modifying the generic name. 
There is no necessary agreement in gender or number. Examples, Clostri- 
dium welchii (Welch's Clostridium.) , Salmonella pidlorum (the Salmonella 
of chicks), Streptococcus lactis (the Streptococcus of milk), Brucella abortus 
(the Brucella of abortion), Clostridium tetani (the Clostridium of tetanus), 
Diplococcus pneumoniae (the Diplococcus of pneumonia). Salmonella 
anatum (the Salmonella of ducks). 

(d) A noun in apposition, that is, an explanatoiy noun. This does not 
agree necessarily with the generic name in gender. This method of naming 
is relativeh^ not common in bacteriology. Examples are Actinomyces 
scabies (the scurf or scab Actinomyces) , Bacillus lacticola (the milk-dweller 
bacillus). Bacillus radicicola (the root-dweller bacillus). 

5. The author of the name is often indicated following the name of the 
species, as Bacillus subtilis Cohn. Sometimes a name is indicated also in 
parenthesis, as Micrococcus luteus (Schroeter) Cohn. This means that 
Schroeter first named the species, giving it the name luteus, but placed it in 
another genus {Bacteridium) . Cohn placed it in a new genus. It should 
be noted that the name of a person, following the name of an organism is 
frequently not the person who first discovered or described it, but the person 
who first gave it the name used. For example, Clostridium welchii (Aligula) 
Holland was first described by Dr. Wm. H. Welch, but not named by him. 
It was named by Migula in honor of Dr. Welch and later it was placed in 
the genus Clostridium by Holland. 

6. Sometimes species of bacteria are subdivided into varieties. These 
are likewise given Latin designations, and the entire name written as: 



44 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Streptococcus lactis var. maltigenes (the Streptococcus of milk producing 
malt flavor). 

Some principles of taxonomy. It is important further that the student 
of bacteriology recognize the meaning of certain terms used regularly in 
classifications. 

(1) Species (plural species). A species of plant (or animal) is assumed 
above to be one kind of plant. But how much difference must exist be- 
tween two cultures of bacteria before one is justified in regarding the organ- 
isms in them as being of distinct kinds or species? No rule can be laid 
down. It depends largely upon convenience and a more or less arbitrary 
decision. As stated by Hitchcock (Descriptive Systematic Botany, New 
York, 1925, p. 8): "The unit of classification is a coherent group of like 
individuals, called a species. The term is difficult to define with precision 
because a species is not a definite entity, but a taxonomic concept." Huck- 
er and Pederson (New York Agric. Exper. Sta. Tech. Bull. 167, 1930, p. 39) 
state: "The difficulty met with among these lower forms in dividing them 
into well-defined groups has led man}'' to question whether these small 
groups or 'species' are natural groups and whether such groups can be 
considered to be similar to 'species' among higher forms. However this 
may be, it is necessary to arrange bacteria as well as possible into groups or 
so-called 'species' for convenience in classification," and again (Hucker, 
New York Agric. Exper. Sta. Tech. Bull. 100, 1924, 29), "characters 
applicable to the differentiation of species must evidence a certain amount 
of constancy when studied over a large series of tests. Furthermore, 
characters adapted to the differentiation of larger natural groups or genera 
should, in addition to constancy, show some correlation with other constant 
characteristics. The presence of this relationship or correlation between 
characters for the division of genera indicates that the groupings are being 
made along natural rather than artificial lines." 

Type culture. It is quite evident that when a new species of bacterium 
is described, it must include the particular culture from which the species 
description was made. This original culture is termed the type culture. 
We may develop a definition as follows : — A species of bacterium is the type 
culture or specimen together with all other cultures or specimens regarded 
b}'' an investigator as sufficiently like the type (or sufficiently closely related 
to it) to be grouped with it. It is self-evident that different investigators 
may not draw the same boundaries for a given species. This leads to some 
practical difficulties, but no better definition has been evolved. 

There are certain special cases which require brief discussion. 

(a) How should one designate the different stages in an organism that 
exhibits a growth cycle? There seems to be increasing evidence that certain 
bacteria show cycles in morphology which parallel to some degree those well 



HOW BACTERIA ARE NAMED AND IDENTIFIED 45 

known among the fungi. Such, for example, may well be the rough (R) 
and smooth (S) types described for manj' bacteria, possibly the filterable 
stages noted by many authors, the so-called G types, etc. It is evident that 
an adequate description of any species of bacterium should include a de- 
scription of each of these stages in the cyclical development wherever such 
is proved to exist. In all other cases in botany and in zoology which 
involve growth stages or cycles one stage has been chosen and designated 
as the mature or adult or perfect stage. In ferns, for example, names and 
classifications are based largely upon the sporophytic generation, in insects 
upon the adult or imago, in the rusts upon the stage in which the teleuto- 
spores are produced. There has been no international agreement as to 
what stage should be thus designated for the bacteria. Beyond doubt, it 
would be the stage which is most easily cultured and studied in the la|3ora- 
tory, the stage with which we are best acquainted in the laboratory. It 
might easily happen in bacteria (as it has with fungi) that two different 
stages of the life cycle of single species have been described and named as 
separate species. When the mistake has been discovered, the name given 
to the mature or perfect stage is the one that is accepted. In general the 
descriptions given in the present volume are those which may be regarded 
as belonging to the perfect stage. Unfortunately it is not yet possible 
accurately to group the stages in many of the bacteria that have definite 
growth cycles. 

It is desirable frequently to designate the stage with which one is working. 
This may be done by some conventional sjinbol, as S (smooth tj'pe), G 
(filterable stage), etc. 

(b) How should one designate variants which differ in some minor re- 
spects from the tj'pe, but which do not constitute growth stages? For 
example, the species Bacillus subtilis normally produces endospores. Sup- 
pose that an asporogenous race is derived from such, agreeing with the 
parent culture in all respects, but showing no tendency to revert to spore 
production. What such an organism should be called is a matter of judg- 
ment. It might frequently be designated as an asporogenous strain, or 
more technically if one desires as a variety. It might be termed, for 
example, Bacillus subtilis var. asporus. In other cases such expressions as 
Diplococcus pneumoniae Type 1, or the Rawlings strain of the typhoid bacil- 
lus may be used. 

Unfortunately there is no general agreement upon the exact significance 
which the word "strain" should have in bacteriology. It is recommended 
that it refer merely to source, e.g. the Rawlings strain of Eberthella typhosa, 
and that it be never used to connote a biological character. This would 
not prevent such expressions as "a non-motile strain of Salmonella suipes- 
tifer". but it would make erroneous a statement to the effect that the A 



46 MANUAL OF DETERMINATIVE BACTERIOLOGY 

strain of influenza vims differs from the B strain in certain ways. In 
other words, "strain" is not a synonym of "type" or "variety". We may 
have as many yellow strains of the typhoid bacillus as we have of cultures 
of it, from different sources or specimens. 

(2) Genus (plural genera). A genus is a group of related species. In 
some cases a genus may include only a single species (is said to be monotypic) 
in most cases several to many species are included in a genus. The ques- 
tion asked above may be paraphrased. How close must be the resemblances 
(how close the relationships) among the species of a group to entitle them 
to inclusion in the same genus? In other words, how is it possible to de- 
limit accurately the boundaries of a genus? This is a matter on which 
there is no agreement, and probably can be none. Much of the confusion 
in modern bacteriological terminology is to be attributed to this fact. 
Nevertheless, in course of time experience tends to delimit many genera 
with reasonable accuracy. As stated by Hitchcock (Descriptive Sys- 
tematic Botany, New York, 1925, p. 9): "Convenience may play a role in 
determining generic lines. Extremely large groups may be broken up on 
the basis of differences of smaller degree not common to a group of closely 
allied species, than if the group consisted of a few species. In general, the 
botanist, in delimiting genera, keeps in mind two important requirements, 
that of showing natural affinities and that of aiding correct identification." 
However, a genus may be defined helpfully in another way. One of the 
species described as belonging to a genus is designated as the type species. 
A genus may be defined then, as including this type species together with 
such other species as the investigator (or taxonomist) regards as sufficiently 
closely related. It is apparent that some authors may draw the lines 
narrowly, others broadly. Some authors, for example, recognize only two 
genera of rod-shaped Ijacteria, one for those without endospores {Bac- 
terium), and one for those producing endospores (Bacillus). These genera 
thus defined are very large, each containing hundreds, perhaps thousands, 
of species. Other students break up these large genera into many smaller 
ones. There is not much point to the question as to which is right and 
which is wrong. A better question is, which is the more convenient, better 
represents relationships, better facilitates diagnosis and proves most useful. 
As organisms become better known, it may be possible through the agency 
of the International Committee on Nomenclature to reach agreements 
where lack of agreement leads to serious confusion or misunderstanding. 

(3) Family. A family in taxonomy is a group of related genera. In 
general the name of the family is made from the name or former name of 
one of their genera by affixing the suffix -aceae to the root. The word is 
regarded as plural. Among bacterial families commonly recognized are 
Bacillaceae, Bacteriaceae, Micrococcaceae, Spirochaetaceae, Actinomycetaceae 



HOW BACTERIA ARE NAMED AND IDENTIFIED 47 

(4) Order. An order is a group of related families. It is named usually 
(not always) by substituting the suffix -ales for -aceae in the name of the 
type family. Among ordinal names that have been used in bacteriology 
are Actinomycetalcs, Spirochaetales, Euhaderiales. 

(5) Class. A class is a group of related orders. In this treatise it is 
considered that the bacteria constitute a class of the plant kingdom, and 
this is named ScMzoniycetes. 

(6) Other categories. Other categories or ra7iks of names are used for 
higher groups. Sometimes families are divided into sub-families, these 
into tribes, these into sulitribes, and these finalh^ into genera. 

How to identify an organism by name. One of the main purposes of a 
manual of determinative bacteriology is to facilitate the finding of the 
correct scientific name of a bacterium. Such is the purpose of this volume. 
It is well, however, to note some of the reasons why this result, the identifi- 
cation of an unknown culture, may not eventuate. Among these reasons 
the following may be listed : 

(1) The unknown organism awaiting identification bj' the investigator 
may easily be one which has never been named, or perhaps adequately 
described. For the most part there has been little effort on the part of 
bacteriologists to describe or name bacteria except as they have been found 
to have some economic significance or possess some striking or unusual 
characteristics. It is quite probable that there are manj^ times as many 
species of bacteria undescribed and named as have been described. Such 
undescribed species are all about us. It is not surprising, therefore, if one 
frequently encounters undescribed species. When such unnamed species 
are encountered, particularly if they are of economic importance or are 
related to such forms, it is highly desirable that they should be described, 
named and the results published and made accessible. 

(2) The unknown organism may have been described and named in 
some publication, but the description and name have been over-looked in 
the preparation of the AIaxual. Perhaps the description has been so 
inadequate or incomplete that it has not been possible to place it in the 
classification. It should be noted that the number of species that have 
been described is so great that no one individual can know them all. 
Progress in classification comes about largely as the result of the work of 
specialists in particular groups. For example. Ford made a studj- of all 
of the aerobic spore-bearing bacteria which he had secured from various 
sources. He studied also the descriptions of such bacteria in the literature, 
and then monographed the group. Similar studies on other groups have 
resulted in more or less complete monographs. Such, for example, are the 
monographs on the intestinal group by Welden and Levine, of the acetic 
bacteria by Hoyer, and Visser 't Hooft, of the cocci b}^ Hucker, of the 



48 MANUAL OF DETERMINATIVE BACTERIOLOGY 

pathogenic spore-bearing anaerobes by the English Commission, by Wein- 
berg, and by others, of the red, rod-shaped bacteria by Hefferan and by 
Breed, of the actinomycetes by Waksman and by Lieske, of the root nodule 
bacteria of legumes by Fred and his co-workers, etc. Unfortunately most 
groups of bacteria have not thus been monographed. It is evidently the 
function of a manual such as this to draw largely upon the work of the 
monographers, and to supplement their achievements as far as possible by 
less satisfactory consideration of the unmonographed groups. 

It is clear that because an organism cannot be identified from this text 
is not proof that it has not been described and named. The species most 
closely related may be determined, then the literature searched carefully 
for species described still more closely related or perhaps one identical. 

(3) It is possible, of course, that an error has been made in the selection 
of the correct name. It is desirable that users of these keys and descrip- 
tions should be familiar with the rules governing the correct choice of names, 
and make suitable corrections where needed. 

Steps in determining the name of an organism. The steps in the identifi- 
cation of an uiikno\\Ti organism are usually the following : 

(1) Preparation of an adequate description of the organism. 

(2) Knowledge of construction and use of keys. 

(3) Determination of order, family and genus by use of key. 
Preparation of description of organism. Before attempting to determine 

the name of an ''unknown" organism an adequate description is essential. 
Just what characteristics must be emphasized depends upon the group in 
which the organism falls. It is desirable that the knowledge of the charac- 
ters of the unknown be as complete as possible. 

Use and construction of keys. An exceptionally clear and satisfactory 
discussion of the making and use of keys and synopses is given by Hitch- 
cock (Descriptive Systematic Botanj^ New York, 1925, p. 104). Anyone 
planning to monograph a group is advised to read this. He states: "A key 
is an orderly arrangement of a series of contrasting or directly comparable 
statements, by which groups of the same category may be distinguished 
and indicated or identified," and "A key is primarily a mechanical device 
by which one may arrive at the name of the ultimate member of the group." 
In general the keys used in this Manual are dichotomous, that is, the suc- 
cessive divisions are in twos, differentiation being into two contrasted 
groups. 

Determination of order, family and genus hyuse of keys. The method of 
doing this is discussed in the introduction beginning on page 1 . 



RULES OF NOMENCLATURE* 

In Paris in 1930, the First International ^Microbiological Congress voted 
to follow the rules of nomenclature agreed upon by International Congresses 
of Botany and Zoology "in so far as they yyiay he applicable and appropriate." 
The adoption of the date of the publication of Species Plantarum by Lin- 
naeus in 1753 as the point of departure for bacteriological nomenclature 
was recommended. This recommendation was approved by the plenary- 
session of the IMicrobiological Congress (Proc. P'' Cong. Internat. Micro- 
biol., Paris, 1930, 3, 1932, 519) and by the plenary session of the Botanical 
Congress (Rept. Proc. 5th Internat. Bot. Cong., 1930, Cambridge, 1931, 
p. 16 and 28). 

This Congress also provided for the organization of an International 
Committee on Bacteriological Nomenclature with two permanent sec- 
retaries : 

1. To represent primarily medical and veterinary bacteriology, — Dr. 

R. St. John-Brooks, Lister Institute, London, England. 

2. To represent primarily other phases of bacteriology', — Dr. R. S. Breed, 

Experiment Station, Geneva, New York, U. S. A. 

During the years that have elapsed since its appointment, this Commit- 
tee has organized and has taken various actions in the interest of a more 
stable nomenclature and classification. Some of these have been completed 
and accepted by the Second International Congress of IMicrobiolog}' held 
in London, 1936. These completed actions are quoted below, and are 
incorporated into the classification used in the descriptive portion of the 
Manual. 

The International Rules of Botanical Nomenclature were originally 
adopted by the International Botanical Congresses of Vienna (1903) and 
Brussels (1910). They were modified by the Cambridge Congress (1930) 
so as to accept the type method, and validate species descriptions of bacteria 
unaccompanied by a Latin diagnosis. Some further but less important 
modifications were made at the Amsterdam Congress (1935) (See Sprague, 
Science, 83, 1936, 416). 

The following are the most important of the rules that are of interest to 
bacteriologists taken from the latest available edition of the Botanical Code 
(Gustav Fischer, Jena, 1935). Sections that were newly adopted or 
amended by the Amsterdam Botanical Congress (1935) are indicated 
in the text. 

* Contributed by Prof. R. S. Breed, New York State Experiment Station, Geneva, 
New York, September, 1938; revised, October, 1943. 

49 



50 MANUAL OF DETERMINATIVE BACTERIOLOGY 

INTERNATIONAL RULES OF BOTANICAL 
NOMENCLATURE, 1930-1935 

Chapter I. General Considerations and Guiding Principles (Art. 1-9) 

Art. 1. Botany cannot make satisfactory progress without a precise system of 
nomenclature, which is used by the great majority of botanists in all countries. 

Art. 2. The precepts on which this precise system of botanical nomenclature is 
based are divided into principles, rules, and recommendations . The principles (Art. 
1-9, 10-14, 15-19) form the basis of the rules and recommendations. The object of the 
rules (Art. 19-74) is to put the nomenclature of the past into order and to provide for 
that of the future. They are always retroactive; names or forms of nomenclature 
contrary to a rule (illegitimate names or forms) cannot be maintained. The recom- 
mendations deal with subsidiary points, their object being to bring about greater 
uniformity and clearness in future nomenclature : names or forms contrary to a recom- 
mendation cannot on that account be rejected, but they are not examples to be 
followed. 

Art. 3. The rules of nomenclature should be simple and founded on considerations 
sufficiently clear and forcible for everyone to comprehend and be disposed to accept. 

Art. 4. The essential points in nomenclature are: (1) to aim at fixity of names; 
(2) to avoid or to reject the use of forms and names which may cause error or ambigu- 
ity or throw science into confusion. 

Next in importance is the avoidance of all useless creation of names. 

Other considerations, such as absolute grammatical correctness, regularity or 
euphony of names, more or less prevailing custom, regard for persons, etc., notwith- 
standing their undeniable importance, are relatively accessory. 

Art. 5. In the absence of a relevant rule, or where the consequences of rules are 
doubtful, established custom must be followed. 

Art. 7. Scientific names of all groups are usually taken from Latin or Greek. 
When taken from any language other than Latin, or formed in an arbitrary manner, 
they are treated as if they were Latin. Latin terminations should be used so far as 
possible for new names. 

Art. 8. Nomenclature deals with: (1) the terms which denote the rank of tax- 
onomic groups (Art. 10-14) ; (2) the names which are applied to the individual groups 
(Art. 15-72). 

Art. 9. The rules and recommendations of botanical nomenclature apply to all 
groups of the plant kingdom, recent and fossil, with certain distinctly specified ex- 
ceptions. 

Chapter II. Categories of Taxonomic Groups, and the Terms 
Denoting Them {Art. 10-14, Rec. I, II) 

Art. 10. Every individual plant belongs to a species (species), every species to a 
genus {genus), every genus to a family (Jamilia), every family to an order (orda) 
every order to a class (classis), every class to a division (diviso). 

Chapter III. N^ames of Taxonomic Groups (Art. 15-72, Rec. III-L) 
Section 1. General Principles: Priority {Art. 15-17, Rec. Ill) 

Art. 15. The purpose of giving a name to a taxonomic group is not to indicate the 
characters or the history of the group, but to supply a means of referring to it. 

Art. 16. Each group with a given circumscription, position, and rank can bear 
only one valid name, the earliest that is in accordance with the Rules of No- 
menclature. 



RULES OF NOMENCLATURE 51 

Section 2. The Type Method {Arl. 18, Rec. IV-VII) 

Art. 18. The application of names of taxonomic groups is determined by means of 
nomenclatural types. A nomenclatural type is that constituent element of a group 
to which the name of the group is permanently attached, whether as an accepted name 
or as a synonym. The name of a group must be changed if the type of that name is 
excluded (see Art. 66). 

Section S. Limitation of the Principle of Priority: Publication, 
Starting-points, Conservation of Names {Art. 19-22) 

Art. 19". A name of a taxonomic group has no status under the Rules, and has no 
claim to recognition by botanists, unless it is validly published (see Art. 37). 

Art. 20. Legitimate botanical nomenclature begins for the different groups of 
plants at the following dates : 

{h) Myxomycetes, 1753 (Linnaeus, Species Plantarum. ed. 1).* 

Art. 21. However, to avoid disadvantageous changes in the nomenclature of 
genera by the strict application of the Rules of Nomenclature, and especially of the 
principle of priority in starting from the dates given in Art. 20, the Rules provide a 
list of names which must be retained as exceptions. These names are by preference 
those which have come into general use in the fiftj' years following their publication, 
or which have been used in monographs and important floristic works up to the year 
1890. 

Section 4. Nomenclature of the Taxonomic Groups According to Their 
Categories {Art. 23-35, Rec. VII-XX) 

1. Names of Groups above the Rank of Family. 

Rec. IX. Orders are designated preferably by the name of one of their principal 
families with the ending -ales. 

2. Names of Families and Subfamilies, Tribes, and Sub-tribes . 

Art. 23. Names of families are taken from the name or former name of one of their 
genera and end in -aceae. 

Art. 24. Names of subfamilies {subfamiliae) are taken from the name of one of the 
genera in the group, with the ending -oideae, similarly for tribes {tribus), with the 
ending -eae, and for subtribes {subtrihus) with the ending -inae. 

3. Names of Genera and Subdivisions of Genera. 

Art. 25. Names of genera are substantives (or adjectives used as substantives), 
in the singular number and written with an initial capital, which may be compared 
with our family names. These names may be taken from any source whatever, and 
may even be composed in an absolute!}' arbitrary manner. 

Recommendation X. Botanists who are forming generic names show judgment and 
taste by attending to the following recommendations: 

(a) Not to make names long or difficult to pronounce. 

(b) Not to dedicate genera to persons quite unconnected with botany or at least 

with natural science, nor to persons quite unknown, 

(c) Not to take names from barbarous languages, unless those names are fre- 

quently cited in books of travel, and have an agreeable form that is readily 
adaptable to the Latin tongue and to the tongues of civilized countries. 

* See page 48 for action on date for Schizomycetes. 



52 MANUAL OF DETERMINATIVE BACTERIOLOGY 

(d) To indicate, if possible, by the formation or ending of the name the affinities 

or analogies of the genus 

(e) To avoid adjectives used as nouns. 

(f ) Not to give a genus a name whose form is rather that of a subgenus or section 

(e.g. Eusideroxylon, a name given to a genus of Lauraceae. This, however, 
being legitimate, cannot be altered). 

(g) Not to make names by combining words from different languages {nomina 

hyhrida). 

4. Names of Species (binary names). 

Art. 27. Names of species are binary combinations consisting of the name of the 
genus followed by a single specific epithet. If an epithet consists of two or more 
words, these must either be united into one or joined by a hyphen. Symbols forming 
part of specific epithets proposed by Linnaeus must be transcribed. 

The specific epithet, when adjectival in form and not used as a substantive, agrees 
with the generic names. 

Recommen dalions : 

XIII. The specific epithet should, in general, give some indication of the appear- 
ance, the characters, the origin, the history or the properties of the species. If taken 
from the name of a person it usually recalls the name of the one who discovered or de- 
scribed it, or was in some way concerned with it. 

XIV. Names of men and women, and also of countries and localities used as spe- 
cific epithets, may be substantives in the genitive (Clusii, saharae) or adjectives 
(Clusiamis, dahitricus) . It will be well, in the future, to avoid the use of the genitive 
and the adjectival form of the same epithet to designate two different species of the 
same genus: e.g. Lysimachia Hemsleyana Maximum. (1891), and L. Hci tsleyi Franch. 
(1895). 

XV. In forming specific epithets botanists will do well to have regard also to the 
following recommendations: 

(a) To avoid those which are very long and difficult to pronounce. 

(b) To avoid those which express a character common to all, or nearly all, the 

species of a genus. 

(c) To avoid using the names of little-known or very restricted localities, unless 

the species is quite local. 

(d) To avoid, in the same genus, epithets which are very much alike, especially 

those which differ only in their last letters. 

(e) Not to adopt unpublished names found in travellers' notes or in herbaria, 

attributing them to their authors, unless these have approved publication. 

(f) Not to name a species after a person who has neither discovered, nor de- 

scribed, nor figured, nor in any way studied it. 

(g) To avoid epithets which have been used before in any closely-allied genus, 
(h) To avoid specific epithets formed of two or more (hyphened) words. 

(i) To avoid epithets which have the same meaning as the generic name 
(pleonasm). 

Section 5. Conditions of Effective Publication (Art. 36) 

Art. 36. Publication is effected, under these Rules, either by sale or distribution 
of printed matter or indelible autographs to the general public, or to specified repre- 
sentative botanical institutions. 

No other kind of publication is accepted as effective : communication of new names 
at a public meeting, or the placing of names in collections or gardens open to the pub- 
lic, does not constitute effective publication. 



RULES OF NOMENCLATURE 53 

Section 6. Conditions and Dales of Valid Publication of 
Names {Art. 37-45, Rcc. XXI-XXIX) 

Art. 37. A name of a taxonomic group is not validly published unless it is both (1) 
effectively published (see Art. 36), and (2) accompanied by a description of the group 
or by a reference to a previously and effectively published description of it. 

Art. 38. From January- 1, 1935, names of new groups of recent plants, the Bacteria 
excepted, are considered as validly published only when they are accompanied by a 
Latin diagnosis. 

Art. 40. A name of a taxonomic group is not validh' published when it is merely 
cited as a synonjTn. 

Art. 42. A name of a genus is not validly published unless it is accompanied (1) 
by a description of the genus, or (2) by the citation of a previously and effectively 
published description of the genus under another name, or (3) by a reference to a 
previously and effectively published description of the genus as a subgenus, section 
or other subdivision of a genus. 

Art. 43. The name of a monotypic new genus based on a new species is validated 
(1) by the provision of a combined generic and specific description, (2) by the pro- 
vision of a plate with analyses showing essential characters; but this applies only to 
plates and generic names published before January 1, 1908. 

Art. 44. The name of a species or of a subdivision of a species is not validly pub- 
lished unless it is accompanied (1) by a description of the group, or (2) by the citation 
of a previously and eflfectiveh' published description of the group under another name, 
or (3) by a plate or figure with analyses showing essential characters; but this applies 
only to plates or figures published before January 1, 1908. 

Art. 45. The date of a name or of an epithet is that of its valid publication (see 
Art. 19, 36). For purposes of priority, however, only legitimate names and epithets 
published in legitimate combinations are taken into consideration (see Art. 60). In 
the absence of proof to the contrary, the date given in the work containing the name 
or epithet must be regarded as correct. 

Botanists will do well in publishing to conform to the following recommendations: 

XXI. Xot to publish a new name without clearly indicating whether it is the 
name of a family or a tribe, a genus or a section, a species or a variety; briefly, with- 
out expressing an opinion as to the rank of the group to which the name is given. 

Xot to publish the name of a new group without indicating its type (see Recom- 
mendation IV). 

XXII. To avoid publishing or mentioning in their publications unpublished 
names which they do not accept, especially if the persons responsible for these names 
have not formally authorized their publication (see Recommendation XV (e). 

XXVI. To give the etymology of new generic names and also of new epithets 
when the meaning of these is not obvious. 

XXVII. To indicate precisely the date of publication of their works and that of 
the placing on sale or the distribution of named and numbered plants when these are 
accompanied by printed diagnoses. In the case of a work appearing in parts, the 
last published sheet of the volume should indicate the precise dates at which the 
different fascicles or parts of the volumes were published as well as the number of 
pages in each. 

XXVIII. When works are published in periodicals, to require the publisher to 
indicate on the separate copies the date (year and month) of publication and also the 
title of the periodical from which the work is extracted. 



54 MANUAL OF DETERMINATIVE BACTERIOLOGY 

XXIX. Separate copies should always bear the pagination of the periodical of 
which they form a part; if desired they may also bear a special pagination. 

Section 7. Citation of Authors' Names for Purposes of 
Precision {Art. 46-49, Rec. XXX-XXXII) 

Art. 46. For the indication of the name (unitary, binary, or ternary) of a group 
to be accurate and complete, and in order that the date may be readily verified it is 
necessary to cite the author who first published the name in question. 

Art. 47. An alteration of the diagnostic characters or of the circumscription of a 
group does not warrant the citation of an author other than the one who first pub- 
lished its name. 

When the changes have been considerable, an indication of their nature and of this 
author responsible for the change is added, the words mutatis charact., or pro parte, 
or excl. gen., excl. sp., excl. var., or some other abridged indication being employed. 

Art. 48. When a name of a taxonomic group has been proposed but not published 
by one author, and is subsequently validly published and ascribed to him (or her) 
by another author who supplied the description, the name of the latter author must 
be appended to the citation with the connecting word "ex." 

If it is desirable or necessary to abbreviate such a citation, the name of the publish- 
ing author, being the more important, must be retained. 

When a name and description by one author are published by another author, the 
word apud is used to connect the names of the two authors, except where the name of 
the second author forms part of the title of a book or periodical in which case the con- 
necting word in is used instead. 

Art. 49. When a genus or a group of lower rank is altered in rank but retains its 
name or epithet, the original author must be cited in parenthesis, followed by the 
name of the author who effected the alteration. The same holds when a subdivision 
of a genus, a species, or a group of lower rank is transferred to another genus or species 
with or without alteration of rank. 

Section 8. Retention of Names or Epithets of Groups which 
are Remodelled or Divided {Art. 60-52) 

Art. 50. An alteration of the diagnostic characters, or of the circumscription of a 
group, does not warrant a change in its name, except in so far as this may be neces- 
sitated (1) by transference of the group (Art. 53-55), or (2) by its union with another 
group of the same rank (Art. 56-57), or (3) by a change of its rank (Art. 58). 

Art. 51. When a genus is divided into two or more genera, the generic name must 
be retained for one of them, or (if it has not been retained) must be re-established. 
When a particular species was originally designated as the type, the generic name 
must be retained for the genus including that species. When no type was designated, 
a type must be chosen according to the regulations which will be given (Appendix I).* 

Art. 52. When a species is divided into two or more species, the specific epithet 
must be retained for one of them, or (if it has not been retained) must be re-estab- 
lished. When a particular specimen was originally designated as the type, the spe- 
cific epithet must be retained for the species including that specimen. When no type 
was designated, a type must be chosen according to the regulations to be given (Ap- 
pendix I). 

* Appendix I has not been published as yet. See Type Basis Code, p. 61. 



RULES OP NOMENCLATUEE 55 

Section 9. Retention of Names or Epithets of Groups Below the Rank 
of Genus on Transference to Another Genus or Species (Art. 5S-55) 

Art. 53. When a subdivision of a genus is transferred to another genus (or placed 
under another generic name for the same genus) without change of rank, its subdivi- 
sional name must be retained, or (if it has not been retained) must be re-established 
unless one of the following obstacles exists: (1) that the resulting association of names 
has been previously published validly for a different subdivision, or (2) that there is 
available an earlier validly published sub-divisional name of the same rank. 

Art. 54. When a species is transferred to another genus (or placed under another 
generic name for the same genus), without change of rank, the specific epithet must 
be retained or (if it has not been retained) must be re-established, unless one of the 
following obstacles exists: (1) that the resulting binary name has been previously and 
validly published for a different species, (2) that there is available an earlier validly 
published specific epithet. 

"When, on transference to another genus, the specific epithet has been applied 
erroneously in its new position to a different plant, the new combination must be 
retained for the plant on which the epithet was originally based, and must be attrib- 
uted to the author who first published it." (Accepted in this revised form at the Am- 
sterdam Botanical Congress, 1935.) 

Art. 55. When a variety or other subdivision of a species is transferred, without 
change of rank, to another genus or species (or placed under another generic or spe- 
cific name for the same genus or species), the original subdivisional epithet must be 
retained or (if it has not been retained) must be re-established, unless one of the 
following obstacles exists: (1) that the resulting ternary combination has been previ- 
ously and validly published for a subdivision based on a different type, even if that 
subdivision is of a different rank; (2) that there is an earlier validly published sub- 
divisional epithet available. 

When the epithet of a subdivision of a species, on transference to another species, 
has been applied erroneouslj' in its new position to a different plant, the epithet must 
be retained for the plant on which the group was originally based. 

Example: The variety micranthum Gren. & Godf. (Fl. France, i, 171 : 1847) of Heli- 
anthemum italicum Pers., when transferred, as a variety to H. penicillatum Thib., 
retains its varietal epithet, becoming H. penicillatum var. micranthum (Gren. & 
Godr.) Grosser (in Engl. Pflanzenreich, Heft 14, 115: 1903). 

Section 10. Choice of Names when Two Groups of the Same Rank are 

United, or in Fungi with a Pleomorphic Life-cycle 

{Art. 56, 57, Rec. XXXIII-XXXV) 

Art. 56. When two or more groups of the same rank are united, the oldest legiti- 
mate name or (in species and their subdivisions) the oldest legitimate epithet is re- 
tained. If the names or epithets are of the same date, the author who unites the 
groups has the right of choosing one of them. The author who first adopts one of 
them, definitely treating another as a synonym or referring it to a subordinate group, 
must be followed. 

Art. 57. Among Fungi with a pleomorphic life-cycle the different successive 
states of the same species {anamorphoses, status) can bear only one generic and spe- 
cific name (binary), that is the earliest which has been given, starting from Fries, 
Systema, or Fries, Synopsis, to the state containing the form which it has been agreed 
to call the perfect form, provided that the name is otherwise in conformity with the 
Rules. The perfect state is that which ends in the ascus stage in the Ascomycetes, 



56 MANUAL OF DETERMINATIVE BACTERIOLOGY 

in the basidium, in the Basidiomycetes, in the teleutospore or its equivalent in the 
Uredinales, and in the spore in the Ustilaginales. 

Generic and specific names given to other states have only a temporary value. 
They cannot replace a generic name already existing and applying to one or more 
species, any one of which contains the "perfect" form. 

The nomenclature of Fungi which have not a pleomorphic life-cycle follows the 
ordinary rules. 

Section 11. Choice of Names when the Rank of a Group is Changed 

Art. 58. When a tribe becomes a family, when a subgenus or section becomes a 
genus, when a subdivision of a species becomes a species, or when the reverse of these 
changes takes place, and in general when a group changes its rank, the earliest legiti- 
mate epithet given to the group in its new rank is valid, unless that name or the result- 
ing association or combination is a later homonym (see Art. 60, 61). 

Section 12. Rejection of Names {Art. 59-69, Rec. XXXVII) 

Art. 59. A name or epithet must not be rejected, changed, or modified merely 
because it is badly chosen, or disagreeable, or because another is preferable or better 
known (see also Art. 69). 

Art. 60. A name must be rejected if it is illegitimate (see Art. 2) . The publication 
of an epithet in an illegitimate combination must not be taken into consideration for 
purposes of priority, "e.xcept as indicated in Art. 61." (Added at the Amsterdam 
Botanical Congress, 1935.) 

A name is illegitimate in the following cases: 

(1) If it was superfluous when published, i.e., if there was a valid name (see Art. 
16) for the group to which it was applied, with its particular circumscription, position 
and rank. 

(2) If it is a binary or ternary name published in contravention of Art. 16, 50, 52, 
or 54, i.e., if its author did not adopt the earliest legitimate epithet available for the 
group with its particular circumscription, position, and rank. 

(3) If it is a later homonym (see Art. 61) (except as regards Art. 54 and 55). 

(4) If it is a generic name which must be rejected under Art. 67. 

(5) If its specific epithet must be rejected under Art. 68. 

Art. 61. A name of a taxonomic group is illegitimate and must be rejected if it is a 
later homonym, that is, if it duplicates a name previously and validly published for a 
group of the same rank based on a different type. Even if the earlier homonj^m is 
illegitimate, or is generally treated as a synonym on taxonomic grounds, the later 
homonym must be rejected. "When an author simultaneously publishes the same 
new name for more than one group, the first author who adopts one of them, or sub- 
stitutes another name for one of them, must be followed." (Added at the Amsterdam 
Botanical Congress, 1935.) 

Art. 62. A name of a taxonomic group must be rejected if, owing to its use with 
different meanings, it becomes a permanent source of confusion or error. A list of 
names to be abandoned for this reason {Nomina ambigua) will form Appendix IV.* 

Art. 63. A name of a taxonomic group must be rejected when its application is 
uncertain {Nomendubium):e.g.,Ervumsoloniense'L. {Cent. II. PI. 28: 1756) is a name 
the application of which is uncertain; it must, therefore, be rejected (see Schinz and 
Thell in Vierteljahrsschr. Nat. Ges. Zurich, viii, 71: 1913). 

* Appendix IV has not been published as yet. 



RULES OF XOMENCLATUEE 57 

Art. 64. A name of a taxonomic group must be rejected if the characters of that 
group were derived from two or more entirely discordant elements, especially if those 
elements were erroneously supposed to form part of the same individual. 

A list of names to be abandoned for this reason (Nomina confusa) will form Ap- 
pendix VI.* 

Art. 65. A name or epithet of a taxonomic group must be rejected when it is based 
on a monstrosity. 

Art. 66. The name of an order, suborder, family or subfamily, tribe or subtribe 
must be changed when it is taken from the name of a genus whicli is known not to be- 
long to the group in question — e.g. if the genus Portulaca were excluded from the 
family now known as Portulacaceae, the residual group could no longer bear the name 
Portulacaceae, and would have to be renamed. 

Art. 67. Names of genera are illegitimate in the following special cases and must 
be rejected: 

(1) When they are merely words not intended as names: e.g. Anonymous Walt. 

(Fl. Carol. 2, 4, 9, etc. : 1788) must be rejected as being a word applied to 28 
different genera by Walter to indicate that they were without names. 

(2) When they coincide with a technical term currently used in morphology un- 

less they were accompanied, when originally published, by specific names 
in accordance with the binary method of Linnaeus. On and after Jan. 1, 
1912, all new generic names coinciding with such technical terms are un- 
conditionally rejected. 

(3) When they are unitary designations of species: e.g. Ehrhart (Phytophylacium: 

1780; and Beitr. iv, 145-150: 1798) proposed unitary names for various 
species known at that time under binary names: e.g. Phaeocephalum for 
Schoeniis fuscus, and Leptostachys for Carex leptoslachys. These names, 
which resemble generic names, should not be confused with them, and 
must be rejected, unless they have been published as generic names by a 
subsequent author. 

(4) When they consist of two words, unless these words were from the first com- 

bined into one, or joined by a hyphen. 
Art. 68. Specific epithets are illegitimate in the following cases and must be re- 
jected: 

(1) When they are merely words not intended as names. 

(2) When they are merely ordinal adjectives being used for enumeration. 

(3) When they exactly repeat the generic name with or without the addition of a 

transcribed symbol. 

(4) When they were published in works in which the Linnean system of binary 

nomenclature for species was not consistently employed. 
Art. 69. In cases foreseen in Art. 60-68 the name or epithet to be rejected is re- 
placed by the oldest legitimate name, or (in a combination) by the oldest legitimate 
epithet. If none exists, a new name or epitliet must be chosen. Where a new epithet 
is required, an author may, if he wishes, adopt an epithet previously given to the 
group in an illegitimate combination, if there is no obstacle to its employment in the 
new position or sense. 

Section 13. Orthography of Names (Art. 70-71, 

Rec. XXXVIII-XLIV) 

Art. 70. The original spelling of a name or epithet must be retained, except in the 

case of a typographic error, or of a clearly unintentional orthographic error. When 

the difference between two generic names lies in the termination, these names must 

* Appendix VI has not been published as yet. 



58 MANUAL OF DETERMINATIVE BACTERIOLOGY 

be regarded as distinct, even though differing by one letter only. This does not apply 
to mere orthographic variants of the same name. 

Note 1. The words "original spelling" in this Article mean the spelling em- 
ployed when the name was validly published. 

2. The use of a wrong connecting vowel or vowels (or the omission of a 

connecting vowel in a specific epithet, or in that of a subdivision of a 
species) is treated as an unintentional orthographic error which may 
be corrected (see Rec. XLIV). "The liberty of correcting a name 
must be used with reserve, especially if the change affects the first 
syllable, and above all the first letter of the name." (Added at the 
Amsterdam Botanical Congress, 1935.) 

3. In deciding whether two or more slightly different names should be 

treated as distinct or as orthographical variants, the essential consid- 
eration is whether they may be confused with one another or not: 
if there is serious risk of confusion, they should be treated as ortho- 
graphic variants. Doubtful cases should be referred to the Execu- 
tive Committee. 

4. Specific and other epithets of Greek origin differing merely by having 

Greek and Latin terminations respectively are orthographic variants. 
Epithets bearing the same meaning and differing only slightly in form 
are (considered as) orthographic variants. The genitive and ad- 
jectival forms of a personal name are, however, treated as different 
epithets (e.g. Lysimachia Hemsleyana and L. Hemsleyi). 
Recommendations : 

XXXVIII. When a new name is derived from a Greek word containing the spiri- 
ius asper (rough breathing), this should be transcribed as the letter h. 

XXXIX. When a new name for a genus, subgenus or section is taken from the 
name of a person, it should be formed in the following manner: — 

(a) When the name of the person ends in a vowel the letter a is added (thus Bou- 

teloua after Boutelou; Ottoa after Otto; Sloanea after Sloane), except when 
the name already ends in a, when ea is added (e.g. Collaea after Colla). 

(b) When the name of the person ends in a consonant, the letters ia are added 

(e.g. Magnusia after Magnus, Ramondia after Ramond), except when the 
name ends in er, when a is added (e.g. Kernera after Kerner). 

(c) The syllables which are not modified by these endings, retain their original 

spelling, even with the consonants k and w or with groupings of vowels 
which were not used in classical Latin. Letters foreign to botanical Latin 
should be transcribed, and diacritic signs suppressed. The Germanic 
o, 6, il become ae, oe, ue; the French e, e, e become generally e. In works 
in which diphthongs are not represented by special type, the diaeresis 
sign should be used where required, e.g., Cephaelis, not Cephaelis. 

(d) Names may be accompanied by a prefix or a suffix, or modified by anagram 

or abbreviation. In these cases they count as different words from the 
original name. 
Examples: Durvillea and Urvillea; Lapeyrousea and Peyrousea; Englera, 
Englerastrum and Englerella; Bouchea and Ubochea; Gerardia and 
Graderia. 
XL. When a new specific or other epithet is taken from the name of a man, it 
should be formed in the following manner: — 

(a) When the name of the person ends in a vowel, the letter i is added (thus Gla- 
zioui from Glaziou, Bureaui from Bureau), except when the name ends in 
a, when e is added (thus balansae from Balansa). 



RULES OF NOMENCLATURE 59 

(b) When the name ends in a consonant, the letters ii are added (thus Magnusii 

from Magnus, Ramondii from Ramond), except when the name ends in 
-er when i is added (thus Kerneri from Kerner) . 

(c) The syllables which are not modified by these endings retain their original 

spelling, even with the consonants korwor with groupings of vowels which 
were not used in classical Latin. Letters foreign to botanical Latin should 
be transcribed and diacritic signs suppressed. The Germanic d, o, u be- 
come ae, oe, ou, the French e, e, e become generally e. The diaeresis sign 
should be used where required. 

(d) When epithets taken from the name of a person have an adjectival form they 

are formed in a similar way (e.g. Geranium Robertianum, Verbena 
Hasslerana) . 
XLI. The same provisions apply to epithets formed from the names of women. 
When these have a substantival form they are given a feminine termination (e.g. 
Cypripedium Hookerae, Rosa Beatricis, Scabiosa Olgae, Omphalodes luciliae). 

XLII. The specific (or other) epithets should be written in conformity with the 
original spelling of the words from which they are derived and in accordance with 
the rules of Latin and latinization. 

Examples: silvestris (not sylvestris) sinensis (not chinensis). 

XLIII. Specific (or other) epithets should be written with a small initial letter, 
except those which are derived from names of persons (substantives or adjectives), or 
are taken from generic "or vernacular" names (substantives or adjectives). 
(Emended Amsterdam Botanical Congress, 1935. See page 61 for actions taken by 
Second International Microbiological Congress, London, 1936 governing Bacteriolog- 
ical Nomenclature.) 

XLIV. In the formation of specific (or other) epithets composed of two or several 
roots taken from Latin or Greek, the vowel placed between the two roots becomes a 
connecting vowel, in Latin i, in Greek o; thus menthifolia, salvifolia, not menthae folia, 
salviafolia. When the second root begins with a vowel and euphony requires, the 
connecting vowel should be eliminated (e.g. lepidantha). The connecting vowels ae 
should be retained only where this is required for etymological reasons (e.g. caricae- 
formis from Carica, in order to avoid confusion with cariciformis from Carex). In 
certain compounds of Greek words no connecting vowel is required, e.g. brachycarpus 
and glycylphyllus . 

Section 14. Gender of Generic Names 
Art. 72. The gender of generic names is governed by the following regulations: — 

(1) "A Greek or a Latin word adopted as a generic name retains its classical 

gender. In cases where the classical gender varies, the author has the 
right of choice between the alternative genders. In doubtful cases, general 
usage should be followed." "The following names, however, whose classi- 
cal gender is masculine, are treated as feminine in accordance with historic 
usage: Adonis, Orchis, Slachys, Diospyros, Strychnos. Hemerocallis (m. in 
Sp. PL: Lat. and Gr. hemercalles n.) is also treated as feminine to bring it 
into conformity with all other generic names ending in is." (Emended 
Amsterdam Botanical Congress, 1935.) See Van Eseltine, Jour. Bact., S6, 
1933, 569, for discussion of the gender of generic names used for bacteria. 

(2) Generic names which are modern compounds formed from two or more Greek 

or Latin words take the gender from the last. If the ending is altered, 
however, the gender will follow it, 

(3) Arbitrarily formed generic names or vernacular names used as generic names 

take the gender assigned to them by their authors. Where the original 



60 MANUAL OF DETERMINATIVE BACTERIOLOGY 

author has failed to indicate the gender, the next subsequent author has 
the right of choice. 

Section 15. Various Recommendations (Rec. XLV-L) 

XLV. When writing in modern languages botanists should use Latin scientific 
names or those immediately derived from them, in preference to names of another 
kind or origin (popular names). They should avoid the use of the latter unless these 
are very clear and in common use. 

XLVII. Only the metric system should be used in botany for reckoning weights 
and measures. The foot, inch, line, pound, ounce, etc., should be rigorously excluded 
from scientific language. 

Altitude, depth, rapidity, etc., should be measured in meters. Fathoms, knots, 
miles, etc., are terms which should disappear from scientific language. 

XLVIII. Very minute dimensions should be reckoned in y. (micromillimeters, 
microns, or thousandths of a millimeter) and not in fractions of millimeters or of 
lines, etc.: fractions encumbered with ciphers and commas easily give rise to mis- 
takes. 

XLIX. Authors should indicate clearly and precisely the scale of the figures which 
they publish. 

L. Temperatures should be expressed in degrees of the centigrade thermometer 
of Celsius. 

Chapter IV. Interpretation and Modification of the Rules (Art. 73, 74) 

Art. 73. A small permanent International Executive Committee is established 
with functions including the following: 

(1) Interpreting the Rules in doubtful cases, and issuing considered "Opinions" 

on the basis of the evidence submitted. 

(2) Considering Nomina conservanda, Nomina ambigua, Nomina dubia and 

Nomina confusa, and making recommendations thereon to the next Inter- 
national Botanical Congress. 

(3) Considering all proposals for the modification of the Rules and reporting 

thereon to the next Congress. 

(4) Reporting on the effects of modifications of the Rules accepted at the pre- 

ceding Congress. 
Art. 74. These Rules can be modified only by competent persons at an Interna- 
tional Botanical Congress convened for the express purpose. Modifications accepted 
at one Congress remain on trial until the next Congress, at which they will receive 
sanction unless undesirable consequences, reported to the Executive Committee, 
show need for further amendment or rejection. 

Eight appendices have been or are to be prepared for this Code as 
follows: (1) fRegulations for determining types, (2) fNomina conservanda 
familiarum, (3) *Nomina generica conservanda, (4) fNomina ambigua, 
(5) fNomina dubia, (6) fNomina confusa, (7) *Representative botanical 
institutions recognized under Art. 34, (8) fNomenclature of garden plants. 

Unfortunately the first appendix which is of greatest interest to bac- 
teriologists has not been prepared. As many bacteriologists, especially 
those in other countries, have not caught the significance of the type species 

* These appendixes have been prepared. 

t These appendixes have not been published as yet. 



RULES OF XOMEXCLATURE 61 

concept as a means of defining bacterial genera, the reader is referred to the 
A\Titings of Hitchcock (Amer. Jour. Bot., 8, 1921, 251; Descriptive Sys- 
tematic Botany, New York, 1925) for an excellent exposition of the value of 
this idea to systematists. 

Hitchcock (1921, p. 252) explains this concept briefly as follows: "The 
old concept was that a genus was a group of species having a given com- 
bination of characters ; a species, similarly, a group of specimens. The new 
type concept is that, from the nomenclatural standpoint, a genus is a group 
of species allied to the type species; a species, a group of individuals similar 
to the type specimen." 

Rules for determining types taken from the Type Basis Code of Nomen- 
clature (Science, 45, 1919, 333; -53, 1921, 312) drawn up by a Committee of 
which Hitchcock was Chairman are quoted as these are the most authori- 
tative rules thus far available. 

Type Basis Code of Nomenclatvre {Hitchcock el al.) 

Article 4. The nomenclatural type species of a genus is the species or one of the 
species included when the genus was originally published. 

If a genus included but one species when originally published, this species is the 
type. 

When more than one species is included in the original publication of the genus, the 
type is determined by the following rules: 

(a) When, in the original publication of a genus, one of the species is definitely 
designated as type, this species shall be accepted as the type regardless of other con- 
siderations. 

If typicus or typus is used as a new specific name for one of the species, this species 
shall be accepted as the type as if it were definitely' designated. 

(b) The publication of a new generic name as an avowed substitute for an earlier 
one does not change the type of the genus. 

(c) If a genus, without an originally designated type, contains among its original 
species one with the generic name used as a specific name, either as a valid name or 
synonym, that species is to be accepted as the type. 

(d) If a genus, when originally published, includes more than one species, and no 
species is definitely designated as type, nor indicated according to (c), the choice of 
the type should accord with the following principles: 

1. Species inquirendae or species doubtfully referred to the genus, or mentioned 

as in any way e.xceptional are to be excluded from consideration in selecting 
the type. 

2. Genera of the first edition of Linnaeus's "Species Plantarum" (1753) 

are usually typified through the citations given in the fifth edition of his 
"Genera Plantarum" (1754) except when inconsistent with the preceding 
articles. 

3. Species which definitely disagree with the generic description (provided others 

agree), or which possess characters stated in the generic description as rare 
or unusual, are to be excluded from consideration in selecting the type. 



62 MANUAL OF DETERMINATIVE BACTERIOLOGY 

RECOMMENDATIONS 

Article 5. In the future it is recommended that authors of generic names definitely 
designate type species; and that in the selection of types of genera previously pub- 
lished, but of which the type would not be indicated by the preceding rules, the fol- 
lowing points be taken into consideration: 

(a) The type species should usually be the species or one of the species which the 
author had chiefly in mind. This is often indicated by 

1. A closer agreement with the generic description. 

2. Certain species being figured (in the same work). 

3. The specific name, such as vulgaris, communis, medicinalis or officinalis. 

(b) The type species should usually be the one best known to the author. It may 
be assumed that an indigenous species (from the standpoint of the author), or an eco- 
nomic species, or one grown in a botanical garden and examined by the author, would 
usually represent an author's idea of a genus. 

(c) In Linnaean genera the type should usually be chosen from those species in- 
cluded in the first technical use of the genus in pre-Linnaean literature. 

(d) The types of genera adopted through citations of non-binomial literature 
(with or without change of name) should usually be selected from those of the original 
species which received names in the first binomial publication. 

(e) The preceding conditions having been met, preference should be shown for a 
species which will retain the generic name in its most widely used sense, or for one 
which belongs to a division of the genus containing a larger number of species, or, 
especially in Linnaean genera, for the historically oldest species. 

(f) Among species equally eligible, the preference should be given to the first 
known to have been designated as the type. 

(g) If it is impossible to select a type under the conditions mentioned above, the 
first of equally eligible species should be chosen. 

While the rules and recommendations of the above botanical codes are ap- 
plicable in general to bacteria and related microorganisms, the fact that 
these are not infallible is evident because the rules developed independently 
by zoologists (see Proc. Biol. Soc. Washington, 39, 1926, 75, for the latest 
Code of Zoological Nomenclature) frequently follow a quite different course. 
In some cases at least the zoological rules will appeal to microbiologists as 
more likely to produce uniformity of usage than the botanical rules. 

For example, microbiologists assembled at the Second International 
Microbiological Congress in London, 1936 accepted (Jour. Bact., S3, 1937, 
445) Art. 13 of the International Rules of Zoological Nomenclature as 
preferable to Rec. 43 of the Botanical Rules to govern bacteriological 
practice. This reads as follows: "While specific substantive names derived 
from names of persons may be written with a capital initial letter, all other 
specific names are to be written with a small initial letter. Some examples 
taken from bacteriological literature are: Salmonella Schottmuelleri or 
Salmonella schottmuelleri. Bacillus Welchii or Bacillus welchii, Acetobacter 
Pasteurianum or Acetobacter pasteurianum, Corynebacterium ovis, Nitro- 
somonas javanensis, Rhizobium japonicum." 



RULES OF NOMENCLATURE 63 

In the Manual all species names are written with a small letter. It is 
felt that the value of a name as a name is lessened if capitals or other marks 
are used to indicate etymology. The derivation of generic and specific 
names is given separately in the descriptive material. 

Likewise for obvious reasons, microbiologists refused (Jour. Bact., 33, 
1937, 445) to follow the botanical and zoological practice which permits the 
use of duplicate generic names, one for an animal and the other for a plant 
group ; and accepted the following rules to govern their practice. 

"a. Generic homonyms are not permitted in the group Protista, 
h. It is advisable to avoid homonyms amongst Protista on the one hand, a plant 
or animal on the other." 

The following actions of the International Committee on Bacteriological 
Nomenclature (Cent. f. Bact., II Abt., 9£, 1935, 481) were confirmed 
(Jour. Bact., 33, 1937, 445). 

Bacillus Cohn 1872 was accepted as a genus conservandum with Bacillus 
subtilis Cohn emend. Prazmowski 1880 as type species. It was agreed 
that Bacillus should be defined so as to exclude bacterial species which do 
not form endospores ; and that the so-called Marburg strain found in type 
culture collections should be accepted as the type or standard strain. 

At the Third International Congress of Microbiology held in New York 
City in September, 1939, a series of recommendations of the Permanent 
International Committees on Bacteriological Nomenclature were accepted 
at the plenary session of the Congress. The third and fourth recommenda- 
tions were: 

3. That the Nomenclature Committee, as at present constituted, shall continue 
to function under the auspices of the International Association of Microbiologists 
as it did under the International Society for Microbiology. 

4. That the International Committee shall select from its membership a Ju- 
dicial Commission consisting of twelve member's, exclusive of members ex officio, 
and shall designate a Chairman from the membership of the Commission. The 
two Permanent Secretaries of the International Committee on Bacteriological 
Nomenclature shall be members ex officio of the Judicial Commission. The Com- 
missioners shall serve in three classes of four commissioners each for nine years, 
so that one class of four commissioners shall retire at every International Con- 
gress. In case of the resignation or death of any Commissioner, his place shall 
be filled for the unexpired term by the International Committee at its next 
meeting. 

By prompt action at and subsequent to the Congress ballots were cast 
in spite of war conditions by 26 of the 62 members of the Permanent Com- 
mittee on Nomenclature. These ballots when examined by the joint Secre- 
taries of the Permanent Committee in November, 1942 were found to have 
resulted in the selection of the persons whose names appear below. These 



64 MANUAL OF DETERMINATIVE BACTERIOLOGY 

are grouped in the three classes specified by the Permanent Committee, 
those receiving the highest number of votes being placed in the nine year 
class, those receiving the next highest in the six year class, etc. Names in 
the classes are arranged alphabetically. 

Elected for nine years. — (The term normally expires in 1948.) R. E. 
Buchanan (U.S.A.), A. J. Kluyver (The Netherlands), E. G. D. Murray 
(Canada), S. Orla Jensen (Denmark): Elected for six years. — (Term nor- 
mally expires in 1945.) J. Howard Brown (U.S.A.), A.-R. Prevot (France), 
J. Ramsbottom (Great Britain), Th. Thjotta (Norway); Elected for three 
years. — (Term normally would have expired in 1942.) A. Lwoff (France), 
R. Renaux (Belgium), A. Sordelli (Argentine), C. Stapp (Germany). 

This announcement was made (Sci., 97, 1943, 370) in the hope that some 
plan for taking tentative action on questions of nomenclature could be 
developed by those members of the Commission who could be reached 
under war conditions. 

While no provision was made in 1939 for the contingencies that have 
arisen, it is felt that those elected should serve until successors are elected. 
Professor R. E. Buchanan has been asked to act as Chairman pro tern of 
the Judicial Commission as there is no possibility of securing an election 
under the rules as adopted. 

Tentative International Rules of Bacteriological Nomenclature were 
presented to the Third International Congress of Microbiology by a U.S.A. - 
Canadian Committee on Compilation of Proposals on Bacteriological Nom- 
enclature. As it proved impossible to give adequate consideration to these 
proposals during the Congress, the following recommendations of the Per- 
manent Committee on Nomenclature were accepted: 

1. That a recognized Bacteriological Code be developed. 

2. That publication of such a proposed Code, when developed, be authorized with 

the proviso that it shall be regarded as wholly tentative, but in the hope that 
it shall be widely tested so that it may be brought up for further considera- 
tion and final disposition at the next Microbiological Congress which should 
normally take place in 1942. 

Copies of this tentative Code have been issued in mimeographed form by 
Prof. R. E. Buchanan, Iowa State College, Ames, Iowa, U.S.A., Chairman 
of the U.S.A. -Canadian Committee and may be obtained from him. 



CLASS SCHIZOMYCETES NAGELI 

(Bericht Verhandl. d. bot. Section d. 33 Versammling deutsch. Naturforsch. u. 
Arzt. Bot. Ztg., 1857, 760.) 

Synonyms: Bacteria Cohn, Beitr. Biol. d. Pflanzen, 1, Heft 1, 1872, 136; Bacteria- 
ceae Cohn, ibid., 237; Bacteriales Clements (as an ordinal name), The Genera of Fungi, 
Minneapolis, 1909, 8; Schizomycetaceae De loni and Trevisan, in Saccardo, Sylloge 
Fungorum, 8, 1889, 923; Schizomycetacea Castellani and Chalmers, Manual of Trop- 
ical Medicine, 3rd ed., 1919, 924; Mychota Enderlein, Eakteriencyclogenie, 1924, 236 
Schizomycetae Stanier and Van Niel, Jour. Bact., 42, 1941, 458. 

Typically unicellular plants. Cells usually small, sometimes ultramicroscopic 
Frequently motile. As in the closely related blue-green algae (Class Schizophyceae) 
the cells lack the definitely organized nucleus found in the cells of higher plants and 
animals. However, bodies containing chromatin which may represent simple nuclei 
are demonstrable in some cases. Individual cells may be spherical; or straight 
curved or spiral rods. These cells may occur in regular or irregular masses or even 
in cysts. Where they remain attached to each other after cell division, they may 
form chains or even definite filaments. The latter may show some differentiation 
into holdfast cells, and into motile or non-motile reproductive cells (conidia). Some 
grow as branching mycelial threads whose diameter is not greater than that of ordi- 
nary bacterial cells, i.e., about one micron. Some species produce pigments. The 
true purple and green bacteria possess pigments much like or related to the true 
chlorophylls of higher plants. These pigments have photosynthetic properties. 
The phycocyanin found in the blue green algae does not occur in the Schizomyceies. 
Multiplication is typically by cell division. Endospores are formed by some species 
included in Eubacteriales. Sporocysts are found in Myxobacteriales. Ultramicro- 
scopic reproductive bodies are found in Borrelomycetaceae. The bacteria are free- 
living, saprophytic, parasitic or even pathogenic. The latter types cause diseases 
of either plants or animals. Seven orders are recognized. 

Key to the Orders and Sub-Orders of the Class Schizomycetes. 

A. Cells rigid, not flexuous. Motility by means of flagella or by a gliding movement. 
1. Cells single, in chains or masses. Not branching and mycelial in character. 
Not arranged in filaments. Not acid-fast. Motility when present by means of 
flagella. 

Order I. Eubacteriales, p. 66. 
a. Do not possess photosynthetic pigments. Cells do not contain free sulfur, 
b. Not attached by a stalk. Do not deposit ferric hydroxide. 

Sub-Order I. Eubacteriineae, p. 67. 
bb. Attached to substrate, usually by a stalk. Some deposit ferric hy- 
» droxide. 

Sub-Order II. Caulobacteriineae, p. 827. 
aa. Possesses photosynthetic chlorophyll -like pigments. Some cells contain 
free sulfur. 

Sub-Order III. Rhodobacteriineae, p. 838. 

65 



66 MANUAL OF DETERMINATIVE BACTERIOLOGY 

2. Organisms forming elongated usually branching and mycelial cells. Multiply 
by cell division, special spores, oidiospores and conidia. Sometimes acid-fast. 
Non-motile. 

Order II. Actinomycetales, p. 895. 

3. Cells in filaments frequently enclosed in a tubular sheath with or without a de- 
posit of ferric hydroxide. Sometimes attached. Motile flagellate and non- 
motile conidia. Filaments sometimes motile with a gliding movement. Cells 
sometimes contain free sulfur. 

Order III. Chlamydobacteriales , p. 981. 
B. Cells flexuous, not rigid. 

1. Cells elongate. Motility, by creeping on substrate. 

Order IV. Myxobacteriales, p. 1005. 

2. Cells spiral. Motility, free swimming by flexion of cells. 

Order V. Spirochaetales, p. 1051. 
Supplements: Groups whose relationships are uncertain. 

1. Obligate intracellular parasites or dependent directly on living cells. 

a. Not ultramicroscopic and only rarely filterable. More than 0.1 micron in 
diameter. 

Group I. Order Rickettsiales , p. 1083. 
aa. Usually ultramicroscopic and filterable. Except for certain pox viruses 
of animals and a few plant viruses, less than 0.1 micron in diameter. 
Group II. Order Virales, p. 1128. 

2. Grow in cell-free culture media with the development of polymorphic struc- 
tures including rings, globules, filaments and minute reproductive bodies (less 
than 0.3 micron in diameter). 

Group III. Family Borrelomycetaceae, p. 1291. 

ORDER I. EUBACTERIALES BUCHANAN. 

(Jour. Bact., g, 1917, 162.) 

Simple and undifferentiated rigid cells which are either spherical or rod-shaped. 
The rods may be short or long, straight or curved or spiral. Some groups or species 
are non-motile, others show locomotion by means of flagella. Elongated cells divide 
by transverse fission and may remain attached to each other in chains. Spherical or- 
ganisms divide either by parallel fission producing chains, or by fission alternating in 
two or three planes producing thus either tetrads or cubes of 8 and multiples of 8 cells. 
Many spherical cells form irregular masses in which the plane of division cannot be 
ascertained. Endospores occur in some species. Some species are chromogenic, 
but only in a few is the pigment photosynthetic (bacteriochlorophyll or other chloro- 
phyll-like pigments). 

A group of rather large, spherical to short rod-shaped, colorless sulfur bacteria » 
which some feel should be included in the order Eubacter tales, has been attached as 
an Appendix to the order Chlamydobacteriales on account of the physiological similar- 
ity between the former organisms and the Beggiatoaceae. These are in Family Achro- 
matiaceae, p. 997. 



KEY TO ORDERS AND SUBORDERS 67 

Sub-Order I. Eubacteriineae Breed, Murray and Kitchens. 
(Jour. Bact., 47, 1944, 421.) 

These are, as the name Eubacteriinae implies, the true bacteria in the narrower 
sense of the word. The cells are rigid and free. Branching occurs only under ab- 
normal conditions of life. They are not attached by holdfasts nor stalks. They 
form no sheaths. One-third of the species form pigments, but these have no photo- 
synthetic properties. Endospores occur in one familj^ {Bacillaceae), rarely in others. 

Key to the Families of the Sub-Order Eubacteriineae. 

I. Xo endospores (except Sporosarcina) . 

A. Can develop on inorganic media. Autotrophic and facultative autotrophic. 

Famil}' I. Nitrobacteriaceae, p. 69. 

B. Cannot develop on inorganic media (exceptions, see Family XII. Bacteriaceae). 
Heterotrophic. 

1. Polar flagellate, straight, curved or spiral rods. Gram-negative. (Some 
species with a single flagellum will be found under Family IV. Rhizobiaceae, 
Family V. Micrococcaceae and Family VIII. Corynebacteriaceae) . 

Family II. Pseudomonadaceae , p. 82. 

2. Large, oval, pleomorphic cells sometimes almost yeast-like in appearance. 
Free living in soil. Fix free nitrogen. Poritrichous flagellation. 

Family III. Azoiobacteriaceae, p. 219. 

3. Peritrichous or non-motile rods, and cocci. 

a. Heterotrophic rods which may not require organic nitrogen for growth. 
Usually motile with one to six or more flagella. Usually form nodules 
or tubercles on roots of plants, or show violet chromogenesis. 
Family IV. Rhizobiaceae, p. 223. 
aa. Heterotrophic rods or cocci which utilize organic nitrogen and usu- 
ally carbohydrates. 

b. Spherical cells in masses, tetrads, and packets. A few species are 
motile with one or two flagella. 
c. Gram-positive to Gram-negative cocci. Not obligate 
parasites. 

Family V. Micrococcaceae, p. 235. 
cc. Gram-negative, and sometimes anaerobic cocci. Obligate 
parasites. 

Family VI. Neisseriaceae, p. 295. 
bb. Spherical cells which grow in pairs and chains; and rods. 

c. Gram-positive cocci and rods. Non-motile (some species of 
Streptococcaceae or Corynebacteriaceae may show motility), 
d. Microaerophilic to anaerobic cocci and rods. Frequently 
in chains. Active in the fermentation of sugars. Never 
reduce nitrates. 

Family VII. Lactobacteriaceae , p. 305. 
dd. Usually aerobic, but sometimes anaerobic rods. Less 
active in the fermentation of sugars. May or may not 
reduce nitrates. 

Family VIII. Corynebacteriaceae , p. 381. 



68 MANUAL OF DETERMIXATIVE BACTERIOLOGY 

oc. Gram-negative rods. When motile, from four to many peri- 
trichous flagella. 

d. Grow well on ordinary media containing peptone. Aerobic 
to facultative anaerobic. 

e. Gram-negative, straight rods which ferment sugars 
with the formation of organic acids, 
f. Produce little or no acid in litmus milk. May or 
may not reduce nitrates. Many yellow chromo- 
gens. Borderline between this and following 
family indistinct. Some species anaerobic. 
Family IX. Achromobacteriaceae, p. 412. 
ff. Produce CO2 and frequently visible gas (CO2 + 
Ho) from glucose. Reduce nitrates. Usually 
from the alimentary, respiratory or urinary tract 
of vertebrates, though some are free-living or even 
plant parasites. 

Family X. Enlerobacteriaceae, p. 443. 
dd. Small Gram-negative rods. Obligate parasites which 
usually require body fluids for growth. Do not grow well 
on ordinary media. Some are anaerobic. 

Family XI. Parvobacteriaceae, p. 545. 
ccc. Rods of varied types not included in above families. 
Aerobic to facultative anaerobic. 

Family XII. Bacteriaceae, p. 596. 
II. Form endospores. Large rods, sometimes in chains. Aerobic to anaerobic. 

Family XIII. Bacillaceae, p. 704. 



*FMIILY I. NITROBACTERIACEAE BUCHANAN 
(Jour. Bact., 2, 1917, 349 and Jour. Bact., 3, 1918, 179.) 

Cells without endospores. Rod-shaped or ellipsoidal except for one spherical spe- 
cies (Xitrosococcus riitrosus). Spiral rods in A'itrosospira and in one species of 
Thiobacillus. Flagella either polar (so far as known), or absent. Gram stain un- 
certain, but presumably Gram-negative for all of the polar flagellate, rod-shaped 
species except for Nitrosomonas monocella which is reported to be Gram-positive. 
Capable of growing without organic compounds, using CO2 as the source of carbon, 
and obtaining their energy by oxidation of ammonia, nitrite, hydrogen, sulfur, or 
thiosulfate. Some species can also utilize organic compounds. Non-parasitic, 
usually soil or water forms. 

Key to the tribes and genera of family Nitrobacteriaceae. 

A. Organisms oxidize ammonia to nitrite, or nitrite to nitrate. Growth on stand- 
ard media very poor or absent. 

Tribe I. Nitrobacterieae, p. 70. 
a. Cells oxidize ammonia to nitrite. 

b. Cells are separate, free or in dense aggregates. Do not form zoogloea. 
c. Cells ellipsoidal. 

Genus I. Nitrosomonas, p. 70. 



cc. Cells spherical, 
ccc. Cells spiral. 



Genus II. A'itrosococcus, p. 71. 
Genus III. Nilrosospira, p. 71. 



bb. Cells form a zoogloea. 

c. The zoogloea is surrounded by a common membrane forming a 
cyst. 

Genus IV. Nitrosocystis, p. 72. 
cc. The massed cells are embedded in slime. No common membrane 
surrounds the cells. 

Genus V. Nitrosogloea, p. 73. 
aa. Cells oxidize nitrite to nitrate. 
b. Cells form no zoogloea. 

Genus VI. Niirobacter, p. 74. 
bb. Cells form a zoogloea. 

Genus VII. Nitrocystis, p. 75. 
B. Organisms oxidize hydrogen. 

Tribe II. Hydrogenomonadeae, p. 76. 
a. Aerobic, non-spore-forming rods with single polar flagellum, or non-motile. 

Genus I. Hydrogenomonas , p. 76. 

* Text revised by Prof. R. S. Breed and Prof. H. J. Conn, Geneva, N. Y., Dec, 
1937. Completely revised by Dr. R. L. Starkey, New Jersey Agricultural Experiment 
Station, New Brunswick, N. J., March, 1943. 

69 



70 MANUAL OF DETERMINATIVE BACTERIOLOGY 

C. Organisms oxidize sulfur or thiosulfate and similar inorganic compounds of 
sulfur. 

Tribe III. Thiobacilleae, p. 78. 
a. Aerobic to anaerobic, non-spore-forming rods with a single polar flagellum 
on each (so far as known), or non-motile. 

Genus I. Thiohacillus, p. 78. 

TRIBE I. NITROBACTERIEAE WINSLOW ET AL. 

(Jour. Bact., 5, 1920, 201.) 

Organisms deriving energy from the o.xidation of ammonia to nitrite or from 
nitrite to nitrate and depend on this oxidation for growth. Fail to grow on media 
containing organic matter in the absence of the specific inorganic materials used as 
sources of energy. Many organic compounds commonly used in standard culture 
media are toxic to this group. 

Genus I. Nitrosomonas Winogradsky. 

{Nitromonas Winogradsky, Ann. Inst. Past., ^, 1890, 257; Arch. Sci. biol., St. Peters' 
burg, 1, 1892, 127; emend. S. and H. Winogradsky, Ann. Inst. Past. 50, 1933, 350.) 

Cells ellipsoidal, non-motile or with a single polar flagellum, occurring singly, in 
pairs, short chains or irregular masses, which are not enclosed in a common membrane. 
Oxidize ammonia to nitrite more rapidly than the other genera of this tribe. From 
Latin, nitrosus, full of soda; M.L. nitrous; and Greek monas, a unit; M.L. a monad. 

The type species is Nitrosomonas europaea Winogradsky. 

1. Nitrosomonas europaea Winograd- Small, compact, sharply defined colo- 

sky. (Arch. Sci. biol., St. Petersburg, nies brownish in color on silica gel. 

1, 1892, 127; Bacterium nitrosomonas Aerobic. 

Lehmann and Neumann, Bakt. Diag., Strictly autotrophic. 

2nd ed., 2, 1899, 187; Pseudomonas Source: Soils of Zurich, Switzerland; 

europaea Migula, in Engler and Prantl, of Gennevilliers, France; and Kazan, 

Die natiirl. Pflanzenfam., 1, la, 1895, 29; Russia. 

Planococcus europaeus Vuillemin, Ann. Habitat: Presumably widely distrib- 

Mycologie, Berlin, 11, 1913, 525.) From uted in soil. 
Latin, europaeus, of Europe. 

Rods: 0.9 to 1.0 by 1.1 to 1.8 microns la. Nitrosomonas europaea var. italica 

occurring singly, rarely in chains of three Perotti (Rendic. d. Accad. d. Lincei 

to four. Possess a single polar flagellum Roma, 15, 1906, 516; Abs. in Cent. f. 

3 to 4 times the length of the rods, or Bakt., II Abt., 19, 1907, 337). Also see 

rarely one at either end. Engel and Skallau (Cent. f. Bakt., II 

Grow readily in aqueous media with- Abt., 97, 305, 1937). 
out organic matter, and containing am- 
monium sulfate, potassium phosphate, 2. Nitrosomonas monocella Nelson, 
and magnesium carbonate. The cells (Cent. f. Bakt., II Abt., 83, 1931, 287.) 
accumulate in soft masses around the From Greek monos, single and Latin 
particles of magnesium carbonate at the cella, room; M.L. single cell, 
bottom of the flask. The liquid is occa- Ovoid rods: 0.6 to 0.9 micron, often 
sionally turbid through development of occurring in pairs. Young cells nearly 
motile swarmer cells or monads. spherical. Motile by means of a single 



FAMILY NITROBACTERIACEAE 



71 



polar flagellum 3 to 5 times as long as the 
rod. Gram-positive (Nelson). Found 
negative by H. J. Conn (personal com- 
munication) . 

No growth in nutrient broth, nutrient 
agar, nutrient or plain gelatin, plain or 
litmus milk, glucose or plain yeast 
water, or on potato. 

Silica gel or agar plates of inorganic 
medium: No typical colonies, but yel- 
lowish brown masses of growth around 
particles of CaCOs in the medium. 

Inorganic liquid medium containing 
ammonium salts: Uniform development 
throughout the liquid as well as in the 
carbonate sediment. 

Even low concentrations of organic 
matter retard or completely inhibit the 
initiation of growth. Plant extracts are 
toxic. 



Free CO2 and O2 necessary for growth. 

Optimum pH 8.0 to 9.0. Poor growth 
below pH 7.0. Some growth above pH 
9.0. 

Optimum temperature for growth and 
oxidation 28°C. 

Aerobic. 

Strictly autotrophic. 

Source: Isolated from field soil. 

Habitat : Presumably widely distrib- 
uted in soil. 

S. Winogradsky and H. Winogradsky 
(Ann. Inst. Pasteur, 50, 1933, 394) have 
described 5 cultures of Nitrosomonas 
which were obtained from soils of France. 
An additional culture has been described 
by H. Winogradsky (Ann. Inst. Pasteur, 
58, 1937, 394) from activated sludge. 



Genus II. Nitrosococcus Winogradsky. 

(Arch. Sci. biol., St. Petersburg, 1, 1892, 127.) 

Cells large spheres, non-motile, not producing zoogloea. Oxidize ammonia to nitrite. 
From Latin, nitrosus, full of soda; and Greek kokkos, grain; M.L. nitrous coccus. 
The type species is Nitrosococcus nitrosus (Migula) Bergey et al. 



1. Nitrosococcus nitrosus (Migula) 
Bergey et al. {Nitrosococcus Winograd- 
sky, Ann. Inst. Pasteur, 6, 1891, 577; 
Arch. Sci. biol., St. Petersburg, 1, 1892, 
127; Micrococcus nitrosus Migula, Syst. 
d. Bakt., ^, 1900, 194 ; Nitrosococcus ameri- 
canus Buchanan, Jour. Bact., 3, 1918, 
180; Manual, 2nd ed., 1925, 35.) From 
Latin, nitrous, full of soda; M.L. nitrous. 

Large spheres, 1.5 to 1.7 microns in 
size, with thick cell membrane. Motil- 
ity could not be demonstrated. Stains 
readily with aniline dyes. Observed 
no zoogloea formation. Gram-positive 



(Omelianski, Cent. f. Bakt., II Abt., 19, 
1907, 263). 

Liquid medium: Turbidity. 

Silica gel : Both dark and light colonies. 
Surface colonies look like small drops of a 
turbid yellowish liquid. 

Aerobic. 

Optimum temperature 20° to 25°C. 

Source : Isolated from soil from Quito, 
Ecuador; Companias, Brazil; Melbourne, 
Australia . 

Habitat : Presumably widely distrib- 
uted in soil. 



Genus III. Nitrosospira Winogradsky. 

(Compt. rend. Acad. Sci., Paris, 192, 1931, 1004; Ann. Inst. Pasteur, 50, 1933, 406.) 

Cells spiral-shaped. Oxidize ammonia to nitrite very slowly. From Latin, 
nitrosus, full of soda; and spira, coil, spiral; M.L. nitrous spiral. 
The type species is Nitrosospira briensis Winogradsky. 



72 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



1. Nitrosospira briensis Winogradsky. 
(Ann. Inst. Pasteur, 50, 1933, 407.) From 
French, Brie, a place name; M.L. of 
Brie. 

Spirals wound tiglitly to form very 
small cylinders as long as 15 to 20 mi- 
crons. Short spirals have the appearance 
of short rods and ellipsoidal cells. Small 
pseudo-cocci were observed in old cul- 
tures. 

Colonies on silica gel : Small colonies 
which occasionally contain cyst-like ag- 
gregates of cells. The cysts are more 
poorly developed than in Nitrosocystis. 

Aerobic. 

Reaction optimum: pH 7.0 to 7.2. 



Source : Uncultivated pasture soil of 
Brie, France. 

Habitat : Presumably widely distrib- 
uted in soil. 

2. Nitrosospira antarctica Winograd- 
sky. (Ann. Inst. Pasteur, 50, 1933, 407.) 
From Greek, antai-kitos , southern, 
antarctic. 

Cells and colonies similar to A'^. brien- 
sis except that the cells are generally 
wound together to form more compact 
spirals. 

Aerobic. 

Reaction optimum: pH 7.0 to 7.2 

Source: Soil from the Antarctic. 

Habitat : Presumably widely distrib- 
uted in soil. 



Genus IV. Nitrosocystis Winogradsky. 
(Compt. rend. Acad. Sci., Paris, 192, 1931, 1003; Ann. Inst. Pasteur, 50, 1933, 399.) 

Cells ellipsoidal or elongated, uniting in compact, rounded aggregates surrounded 
by a common membrane to form cysts. The cysts disintegrate to free the cells, par- 
ticularly when transferred to fresh media. Within the cyst, the cells are embedded 
in slime. Oxidize ammonia to nitrite at a rate intermediate between Nitrosomonas 
and Nitrosospira. From Latin, nitrosus, full of soda; and Greek, kystis, bladder; 
M.L. nitrous cyst. 

The type species is Nitrosocystis javanensis comb. nov. 



1. Nitrosocystis javanensis comb. nov. 
{Nitrosomonas javanensis Winogradsky, 
Arch. Sci. biol., St. Petersburg, 1, 1892, 
127; Pseudomonas javanensis Migula, 
in Engler and Prantl, Die natlirl. 
Pflanzenfam., 1, la, 1895, 30; Compt. 
rend. Acad. Sci., Paris, 192, 1931, 1003.) 
From Latin, of Java. 

Small ellipsoidal cells having a diam- 
eter of 0.5 to 0.6 micron. Possess a polar 
flagellum 20 times as long as the rods. 

In liquid medium produces very com- 
pact zoogloeal masses of cells and motile 
swarmers. The large zoogloea are them- 
selves composed of smaller compact 
aggregates of cells. 

On silica gel the colonies are circular to 
elliptical becoming clear or light brown. 

Aerobic. 

Strictly autotrophic. 

Source: Soil of Buitenzorg, Java; 
Tokyo, Japan; La Reghaia, Tunisia. 



Habitat : Presumably widely distrib- 
uted in soil. 

2. Nitrosocystis coccoides nom. nov. 
(Nitrosocystis a, S. Winogradsky and H. 
Winogradsky, Ann. Inst. Pasteur, 50, 
1933, 401.) From Greek, kokkos, a grain; 
eidos, form, shape; M.L. coccus-like. 

Ellipsoidal cells about 1.5 microns in 
diameter. Occur as compact aggregates 
of cells imbedded in mucus and sur- 
rounded by a thickened capsule to form 
cyst-like bodies. Cells rarely solitary 
but more often in pairs and in small 
groups of four or more. Probably motile. 
The mucus which surrounds the cells is 
not readily stained, whereas the outside 
coating stains more easily. 

Colonies on silica gel : As colonies de- 
velop, the coating of CaCOs on the gel 
becomes yellowish and dissolves and the 



FAMILY NITROBACTERIACEAE 



73 



colony appears as a bulbous, angular, 
brown body which may become 0.5 mm. 
in diameter. The cells are held firmly 
together in these irregularly shaped bul- 
bous aggregates. 

Aerobic. 

Source: Poor soils of Brie and else- 
where in France. 

Habitat : Presumably widely distrib- 
uted in forest and manured soils. 

A similar culture called Nitrosocystis 
B.A. was isolated from activated sludge 
by H. Winogradskj' (Compt. rend. Acad. 
Sci., Paris, 200, 1935, 1888; Ann. Inst. 
Pasteur, 58, 1937, 326). It produced 
compact, bulbous, dented cj'st-like ag- 
gregates of cells having a yellow color. 
The colonies produced clear zones on 
silica gel coated with CaCOs. These 
cysts were composed of oval or elongated 
coccoid cells imbedded in mucus and sur- 
rounded by a thickened capsule, com- 



posed of two layers. The cells become 
dispersed from the cysts as motile cells 
and form new colonies. This culture 
differs from A^. coccoides in that the 
colonies have a pale reddish yellow color 
and the oval cells are 0.5 by 1.5 microns 
in size. 

Cultures of N itrosocystis were obtained 
by Rommell (Svensk. botan. Tidskrift, 
26, 1932, 303) from forest soils. Ivingma 
Boltjes (Arch. f. Mikrobiol., 6, 1935, 79) 
obtained cultures which produced masses 
of cells, some of which were loose and 
others compact. They were not believed 
to be true zoogloea since no capsule or 
slimy substance was noted. The de- 
velopment of true cysts by nitrifying 
bacteria was questioned. Winogradsky 
(Bull. d. I'Inst. Pasteur, 33, 1935, 1074) 
concluded that Kingma Boltjes worked 
with a culture of Nitrosocystis and not of 
NilrosomoJias as was believed. 



Genus V. Nitrosogloea H. Winogradsky. 
(Compt. rend. Acad. Sci., Paris, £00, 1935, 1887; Ann. Inst. Pasteur, 58, 1937, 335.) 

Cells ellipsoidal or rod-shaped. Embedded in slime to form zoogloea. No common 
membrane surrounds the cells aggregates. Oxidize ammonia to nitrite. From Latin, 
nitrosus, full of soda; and Greek, gloea, glue, jelly; M.L. nitrous jelly. 

The type species is Nitrosogloea merismoides H. Winogradsky. 



1. Nitrosogloea merismoides H. Wino- 
gradsky. {Nitrosocystis "/", H. Wino- 
gradsky, Trans. Third Intern. Cong. 
Soil Sci., Oxford, 1, 1935, 139; Compt. 
rend. Acad. Sci., Paris, 200, 1935, 1887; 
Ann. Inst. Pasteur, 58. 1937, 333.) From 
Greek, merismos, a dividing, division; 
eidos, form, shape; M.L. division-like. 

Ellipsoidal cells: 0.5 by 1.5 microns. 
Oval cells or short rods forming tetrads 
or chains, each group with its own sheath. 
The groups vary in shape to produce 
branched chains, irregular or compact 
aggregates. 

Colonies on silica gel : Cells encased in 
a pale yellow mucilage giving the colony 
a dull appearance. Colony surface stud- 
ded with little humps. 



Aerobic. 

Source: Activated sludge. 

Habitat: Unknown. 

2. Nitrosogloea schizobacteroides H. 

Winogradsky. (Nitrosocystis "77", H. 
Winogradsky, Trans. Third Intern. Cong. 
Soil Sci., Oxford, 1, 1935, 139; Compt. 
rend. Acad. Sci., Paris, 200, 1935, 1887; 
Ann. Inst. Pasteur, 58, 1937, 333.) From 
Greek, schizo, to split; bakterion, a small 
rod; eidos, form, shape; M.L. like a divid- 
ing rod. 

Rods: Elongated rods or short fila- 
ments 3 to 4 microns long. 

Colonies on silica gel: Flat groups of 
cells are produced which are united in a 
common sheath. The aggregates form a 



74 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



pseudo-tissue of interwoven filaments 
suggestive of a fungus pad. The pad can 
be removed as a unit from the medium. 
• Aerobic. 

Source: Activated sludge. 

Habitat: Unknown. 

3. Nitrosogloea membranacea H. 

Winogradskjr. (Nitrosocystis "HI", H. 
Winogradsky, Trans. Third Intern. Cong. 
Soil Sci., Oxford, 1, 1935, 139; Compt. 
rend. Acad. Sci., Paris, 200, 1935, 1887; 
Ann. Inst. Pasteur, 58, 1937, 333.) From 



Latin, membranaceus , of skin or 
membrane. 

Ellipsoidal cells conamonly in pairs and 
also solitary. 

Colonies on silica gel : Appear as dull 
mucoid material with a pale straw color. 
The cells are held firmly together so that 
the entire colony is easily picked up with 
the transfer needle. No structural units 
within the colony. 

Aerobic. 

Source: Activated sludge. 

Habitat: Unknown. 



Genus VI. Nitrobacter Winogradsky. 

(Winogradsky, Arch. Sci. biol., St. Petersburg, /, 1892, 127; Nitromonas Orla- 
Jensen, Cent. f. Bakt., II Abt., 22, 1909, 334; not Nitromonas Winogradsky, Ann. 
Inst. Past., 4, 1890, 257; Nitrohacterium Castellani and Chalmers, Manual Trop. Med., 
1919, 933.) 

Cells rod-shaped. Oxidize nitrite to nitrate. From Latin, nitrum, soda; M.L. 
nitre; and Greek baktron, a small rod. 
The type species is Nitrobacter winogradskyi Buchanan. 



1. Nitrobacter winogradskyi Buch- 
anan. {Nitrobacter Winogradsky, Arch. 
Sci. biol., St. Petersburg, 1, 1892, 127; 
Bacterium nitrobacter Lehmann and Neu- 
mann, Bakt. Diag., 2nd ed., 2, 1899, 187; 
Bacillus nitrobacter Lohnis, Vorlesungen 
landw. Bakt., Berlin, 1913, 152; Buch- 
anan, Jour. Bact., 3, 1918, 180; Nitrobac- 
terium nitrobacter Castellani and Chal- 
mers, Manual Trop. Med., 1919, 933.) 
Named for S. Winogradsky, 1856- , the 
Russian microbiologist, who first isolated 
these bacteria. 

Description taken from Gibbs, Soil 
Sci., 8, 1919, 448. 

Short, non-motile rods with gelatinous 
membrane, 0.6 to 0.8 by 1.0 to 1.2 mi- 
crons. Does not stain readily. Gram- 
negative (Omelianski, Cent. f. Bakt., II 
Abt., 19, 1907, 263.) 

Can be cultivated on media free of 
organic matter. Sensitive to certain 
organic compounds. 

Washed agar colonies : In 7 to 10 days 
very small, light brown, circular to ir- 
regular colonies, becoming darker. 



Silica gel : Colonies smaller but more 
dense than on washed agar. 

Washed agar slant: In 7 to 10 days 
scanty, grayish streak. 

Inorganic solution medium: After 10 
days flocculent sediment. Sensitive to 
ammonium salts under alkaline condi- 
tions. 

Nitrite is oxidized to nitrate. 

Aerobic. 

Strictly autotrophic. 

Optimum temperature 25° to 28°C. 

Source: Soil. 

Habitat: Presumably widely distrib- 
uted in soil. 

2. Nitrobacter agile Nelson. (Cent, f . 
Bakt., II Abt., 83, 1931, 287.) From 
Latin agile, quick, agile, motile. 

Rods: 0.5 by 0.8 to 0.9 micron, occur- 
ring singly, sometimes in pairs or larger 
aggregates. Rapidly motile with a long, 
thin, polar flagellum often 7 to 10 times 
as long as the rod. (Non-motile culture 
obtained by Ivingma Boltjes, Arch. f. 
Mikrobiol., 6, 1935, 79.) Gram -negative. 



FAMILY NITROBACTERIACEAE 



75 



No growth in nutrient broth, nutrient 
agar, nutrient or plain gelatin, litmus or 
plain milk, glucose or plain yeast water, 
or on potato. 

Nitrite agar : After two weeks, produces 
semi-spherical, minute, nearly transpar- 
ent colonies. Oxidation usually com- 
plete in 10 to 14 days. 

Inorganic liquid medium containing 
nitrite : Produces uniformly dispersed 
growth. 

Optimum pH 7.6 to 8.6. Limits of 
growth 6.6 to 10.0. 



Temperature relations: Optimum for 
growth 25° to 30°C. Optimum for oxida- 
tion 28°C. No oxidation at 37°C. Ther- 
mal death point 60°C. for five minutes. 

Strictly autotrophic. 

Aerobic. 

Source : Isolated from greenhouse soils 
and from sewage effluents in Madison, 
Wisconsin. 

Habitat: Presumably widely distrib- 
uted in soil. 



Genus VII. Nitrocystis H. Winogradsky. 

(Trans. Third Intern. Cong. Soil Sci., O.xford, i, 1935, 139 ; Nitrogloea H. Winogradsky, 
Comp. rend. Acad. Sci., Paris, 200, 1935, 1888.) 

Cells ellipsoidal or rod-shaped. Embedded in slime and united into compact zoo- 
gloeal aggregates. Oxidize nitrite to nitrate. From Latin, nitrum, soda; M.L. 
nitre; and Greek, kystis, bladder; M.L. nitric cyst. 

The type species is Nitrocystis sarcinoides. 



1. Nitrocystis sarcinoides H. Wino- 
gradsky. {Nitrocystis B. A., Winograd- 
sky, H., Compt. rend. Acad. Sci., Paris, 
200, 1935, 1888; Nitrocystis "/" and 
"//", Winogradsky, H., Trans. Third 
Intern. Cong. Soil Sci., Oxford, 1, 1935, 
139; Ann. Inst. Pasteur, 58, 1937, 336.) 
From Latin, sarcina, a packet; M.L. 
Sarcina, a genus; Greek, eidos, form; 
M.L. Sarcina-like. 

Rods: Small rods 0.5 by 1.0 micron. 
Cells ellipsoidal or wedge-shaped and 
grouped in sarcina-like packets. 

Colonies on silica gel : On the surface of 
gel coated with kaolin the colonies appear 
as small raised amber warts. The colon- 
ies grow up to 5 mm. in diameter. The 
colonies are viscous and sticky when 
young and they become brown with age, 
shrink, and look like scales and become 
hard like grains of sand. Each colony is 
enveloped in several layers of a thick 
slime which holds the cells together so 
that the entire colony can be removed 
with a transfer needle. 

Aerobic. 

Source: Activated sludge. 

Habitat: Unknown. 



2. Nitrocystis micropunctata H. Wino- 
gradsky. {Nitrocystis "III", Wino- 
gradsky, H., Trans. Third Intern. Cong. 
Soil Sci., Oxford, 1, 1935, 139; Nitrogloea 
micropunctata Winogradsky, H., Compt. 
rend. Acad. Sci., Paris, 200, 1935, 1888; 
Ann. Inst. Pasteur, 58, 1937, 326.) From 
Greek, mikros, small, little; and Latin, 
punctatus, spotted; M.L. with small 
spots. 

Cells are ellipsoidal rods about 0.5 
micron in diameter which stain poorly 
except at the ends. Encased in a viscous 
slime. 

Colonies on silica gel : Like N, sar- 
cinoides except that the colonies are more 
clear and they have a more plastic con- 
sistency. The cells are not held together 
by the slime in the colony as with N. 
sarcinoides. The capsule is more readily 
differentiated in old colonies. 

Aerobic. 

Source: Activated sludge. 

Habitat: Unknown. 

Appendix: The following have been 
placed in the Tribe Nitrobacterieae, some- 
times incorrectly so : 

Bactoderma alba Winogradsky. (Aim. 



76 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Inst. Pasteur, 50, 1933, 414.) From soil. 
This is the type species of genus Bnclo- 
derma Winogradsky. 

Bactoderma rosea Winogradsky (loc. 
cii., p. 415). Isolated from soil. 

Bacterium nitrificans Chester. (Ni- 
tratbildner aus Northeim, Burri and 
Stutzer, Cent. f. Bakt., II Abt., 1, 1895, 
735; Chester, Ann. Rept. Del. Col. Agr. 
Exp. Sta., 9, 1897, 94; Bacillus nitrificans 
Chester Man. Determ. Bact., 1901, 239; 
Achromobacter nitrificans Bergey et al., 
Manual, 1st ed., 1923, 137.) From soil. 
Description of this organism was shown 
by Winogradsky (Cent. f. Bakt., II 
Abt., 2, 1896, 415 and 449) to have been 
based on impure cultures. 

Microderma yriinulissima Winogradsky. 
From soil. This is the type species of 
genus Microderma Winogradsky. 

Microderma vacuolata Winogradsky 
(loc. cit.). Isolated from soil. 

Ni'.rosobacillus thermophilus Campbell. 
See Bacillus appendix. 

Nilrobacier flavum Sack. (Cent. f. 
Bakt., II Abt., 62, 1924, 20.) Isolated 
from garden earth. See description. 



Manual, 5th ed., 1939, 74. Heterotrophic 
and does not belong here (Kingma 
Boltjes, Arch. f. Mikrobiol., 6, 1935, 83). 

Nitrobacter oligotrophtm Beijerinck. 
(Folia Microbiol., 3, 1914, 91 ; Verzamelde 
Geschriften van M. W. Beijerinck, 5, 
1922, 190.) Isolated from soil. On cul- 
tivation this species lost its autotrophic 
habit and became heterotrophic. The 
organism was then called Nitrobacter 
polytrophum Beijerinck. 

Nitrobacter opaciim Sack (loc. cit. p. 
21). Source and relationships as above. 
See Manual, 5th ed., 1939, 75. 

Nitrobacter punctatum Sack {loc. cit., 
p. 20). Source and relationships as 
above. See Manual, 5th ed., 1939, 75. 

N itrobacter roseo-album. Sack {loc. cit., 
p. 17; Serratia roseo-alba Bergey et al., 
Manual, 3rd ed., 1930, 125.) Source and 
relationships as above. See description, 
Manual, 5th ed., 1939, 74. 

Nitrosomonas groningensis Sack. 
(Cent. f. Bakt., II Abt., 64, 1925, 34.) 
Source and relationships as above. See 
description, Manual, 5th ed., 1939, 77. 



TRIBE II. HYDROGENOMONADEAE PRIBRAM. 

(Jour. Bact., 18, 1929, 370.) 

Short rods, non-motile or with lophotrichous flagella. Organisms capable of deriv- 
ing energy from oxidation of hydrogen. They probably grow well on organic media 
without hydrogen, although this has not been shown to be true for all species. 

Genus I. Hydrogenomonas Orla-Jensen.* 

(Cent. f. Bakt., II Abt., 22, 1909, 311.) 

As the only genus of the tribe, its definition is identical with the definition of the 
tribe. From Greek hydor, water; genos, producing and monas, a unit. 
The type species is Hydrogenomonas pantoiropha (Kaserer) Orla-Jensen. 

* This group of bacteria is characterized by the ability to grow in substrates contain- 
ing no organic matter and to utilize elemental hydrogen as the source of energy for 
growth. Under these conditions CO2 is used as the source of carbon. Bacteria with 
similar physiological characteristics but differing in morphology are placed in the 
genera Bacterium, Bacillus and Clostridium. Although other bacteria and even cer- 
tain algae have enzyme systems which can activate hydrogen and reduce CO2 in the 
process, there is no evidence that these organisms are able to grow in inorganic media 
with hydrogen as the exclusive source of energy (See : Stephenson and Stickland, Bio- 
chem. .Tour., 25, 1931, 205, 215; Woods, Biochem. .Jour., 30, 1936, 515; Lee and Umbreit, 
Cent. f. Bakt., II Abt., 101, 1940, 354; Gaffron, Amer. Jour. Bot., 27, 1940, 273). 



FAMILY XITROBACTERIACEAE 



77 



Key to the species of genus Hydrogenomonas. 

A. Xot sensitive to high O2 concentrations. Growth in solution media under 

autotrophic conditions characterized by turbidity without pellicle formation. 

1. Hydrogenomonas pantotropha. 

B. Sensitive to high O2 concentrations. Growth in solution media under auto- 

trophic conditions characterized by pellicle adhering to walls of container. 

2. Hydrogenomonas vitrea. 

C. Sensitive to high O2 concentrations. Growth in solution media under auto- 

trophic conditions without pellicle formation. 

3. HTjdrogenomonas flava. 



1. Hydrogenomonas pantotropha 
(Kaserer) Orla-Jensen. [Bacillus pan- 
totrophus Kaserer, Cent. f. Bakt., II 
Abt., 16, 1906, 688; Orla-Jensen, Cent, 
f. Bakt., II Abt., 22, 1909, 311.) From 
Greek pantos, everything and trophos, 
feeds on; M.L. omnivorous. 

Rods: 0.4 to 0.5 by 1.2 to l.o microns 
with rounded ends. Occur singly, in 
pairs, and in chains. Encapsulated. Ac- 
tively motile by means of a single long 
polar flagellum. Gram stain not re- 
corded. Bipolar staining in old cultures. 

Inorganic solution : When cultivated 
under an atmosphere of O2, CO2 and 
Ho, the liquid becomes turbid without 
pellicle formation. 

Inorganic solid media : When cultivated 
under an atmosphere of O2, CO2 and H2, 
the colonies are yellow and slimy, and the 
agar plates have an odor resembling hot 
soapy water. 

Gelatin colonies: Yellow, smooth, 
rarely concentrically ringed or greenish. 

Gelatin stab: Growth only at surface. 
.\s a rule no liquefaction. 

Agar colonies: Same as on gelatin, 
greenish, often slimy. 

Broth: Turbid, somewhat slimy, and 
occasional pellicle. 

Milk: Xo coagulation. A yellow pelli- 
cle forms. Medium becomes slimy and 
assumes a dirty flesh color. 

Potato: Moist, yellow, glistening. 

Indole is not formed. 

Hydrogen sulfide is not formed. 

Nitrite is not produced from nitrate. 

Does not act on carbohydrates. 

Aerobic. 



Optimum temperature 28° to 30°C. 

Facultative autotroph. 

Distinctive characters : Develops auto- 
trophically in inorganic medium under an 
atmosphere of Ho, O2 and CO2. Oxidizes 
hydrogen to water and uses CO2 as the 
source of carbon for growth. 

Source : Isolated from soil near Vienna. 

Habitat : Probably widely distributed 
in soil. 

2. Hydrogenomonas vitrea Xiklewski. 
(Jahrb. f. wissensch. Botanik, 48, 1910, 
113). From Latin vitreus, of glass, trans- 
parent. 

Rods: 2.0 microns in length, cells ad- 
hering to each other as by slime. Mo- 
tility not observed. 

Agar colonies on inorganic medium in 
presence of Ho, Oo and CO2: Delicate, 
transparent, with slight fluorescence, and 
yellow center. Surface folded. Do not 
develop readih' beneath the surface of 
medium. 

Agar streak on inorganic substrate : 
Same as agar colonics except that growth 
is spreading. 

Inorganic liquid medium in presence of 
H2, O2 and COo: Pellicle, adherent to 
wall of tube. Good development when 
there is from 2 to 8 per cent oxygen in 
the gas. At higher Oo concentrations 
good growth occurs only in association 
with H. flava or other bacteria. 

Oxidizes hydrogen to water. 

Microaerophilic, growing in an atmos- 
phere of low oxygen tension, not exceed- 
ing 8 per cent. 

Facultative autotroph. 



78 



MANUAL OF DETERMINATIVE BACTERIOLOGT 



Distinctive characters : Grows in sub- 
strates containing no organic matter and 
produces a pellicle. 

Source : Isolated from mud, garden 
soil, pasture land, vegetable mold, and 
peat. 

Habitat: Presumably widely distrib- 
uted in soil. 

3. Hydrogenomonas flava Niklewski. 
(Jahrb. f. wissensch. Botanik., 48, 1910, 
113; emend. Kluyver and Manten, 
Antonie v. Leuwenhoek, 8, 1942, 71.) 
From Latin flavus, yellow. 

Rods: 1.5 microns in length. Motility 
by polar flagella. Gram-negative. 

Agar colonies on inorganic medium in 
presence of H2, O2 and CO2: Small, 
smooth, yellow, shining, adhering to 
medium. Develop well below surface of 
medium, but growth is paler. 

Gelatin not liquefied. 

Inorganic liquid medium in presence of 
Hj, O2, and CO2: No pellicle formation. 
Good development when there is from 2 
to 8 per cent oxygen in the gas . At higher 
O2 concentrations good growth occurs 



only in association with H. viirea or other 
bacteria. 

Oxidizes hydrogen to water. 

Microaerophilic, growing in an atmos- 
phere of low oxygen tension, not exceed- 
ing 8 per cent. 

Facultative autotroph. 

Distinctive characters : Found singly 
on slides whereas the rod-shaped cells of 
Hydrogenomonas vitrea tend to cling to- 
gether in masses. Colonies on agar 
opaque, not transparent. 

Source : Same as H. vitrea. 

Habitat : Presumably widely distrib- 
uted in soil. 

Appendix : Incompletely described 
species are found in the literature as 
follows : 

Hydrogenomonas agilis Niklewski. 
(Jubliaumsschrift f. Prof. E. Godlewski. 
Kosmos, Lemberg, 1913; See Cent. f. 
Bakt., II Abt., W, 1914, 430.) From 
soil. 

Hydrogenomonas minor Niklewski. 
(Jubliaumsschrift f. Prof. E. Godlewski. 
Kosmos, Lemberg, 1913; See Cent. f. 
Bakt., II Abt., 40, 1914, 431.) From soil. 



TRIBE III. THIOBACILLEAE BERGEY, BREED AND MURRAY. 

(Preprint, IVIanual, 5th ed., Oct., 1938, v.) 

Organisms capable of deriving their energy from oxidation of sulfur or sulfur com- 
pounds. Most species do not grow on organic media. 



Genus I. Thiobacillus Beijerinck. 

(Beijerinck, Cent. f. Bakt., II Abt., 11, 1904, 593; Sulfomonas Orla-Jensen, Cent, f . 
Bakt., II Abt., 22, 1909, 314; not Thiobacillus Ellis, Sulphur bacteria, London, 1932, 
130; Thiobacterium Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 517; not 
Thiobacteri^im Janke, Allgemeine Tech. MikrobioL, 1, 1924, 68, Leipzig.) 

Small Gram-negative, rod-shaped cells. Non-motile or motile by means of a single 
polar flagellum. Derive their energy from the oxidation of incompletely oxidized 
sulfur compounds, principally from elemental sulfur and thiosulfate but in some cases 
also from sulfide, sulfite, and polythionates. The principal product of oxidation is 
sulfate, but sulfur is sometimes formed. They grow under acid or alkaline conditions 
and derive their carbon from carbon dioxide or from bicarbonates in solution; some 
are obligate and some facultative autotrophic. One species is facultative anaerobic. 
From Greek theion, sulfur and Latin bacillus, a small rod. 
The type species is Thiobacillus thioparus Beijerinck. 



FAMILY NITROBACTERIACEAE 



79 



Key to the species of genus Thiobacillus. 

I. Aerobic. 

A. Strictly autotrophic. 

1. Optimum reaction for growth close to neutrality. 

1. Thiobacillus thioparus. 

2. Optimum reaction for growth pH 2.0 to 3.5. 

2. Thiobacillus thiooxidans . 

B. Facultative autotrophic. 

3. Thiobacillus novellus. 

4. Thiobacillus coproliticus . 

II. Anaerobic in presence of nitrate. 

5. Thiobacillus denitrificans. 



1. Thiobacillus thioparus Beijerinck. 
(Cent. f. Bakt., II Abt., 11, 1904, 593; 
Nathanson, Mitt. Zool. Station Neapel, 
15, 1902, 655 ; Sulfomonas thioparus Orla- 
Jensen, Cent. f. Bakt., II Abt., 22, 1909, 
326). From Greek theion, sulfur and 
paro, to make. 

Thin, short rods, 0.5 by 1 to 3.0 mi- 
crons. Motile (non-motile culture re- 
ported. See Starkey, Soil Sci., 39, 1935, 
197.) Gram-negative. 

Thiosulfate medium (liquid) : Pellicle 
consisting of cells and free sulfur. 

Thiosulfate agar: Colonies small, cir- 
cular, whitish yellow due to precipitated 
sulfur. 

Optimum reaction : Close to neutrality. 

Strictly autotrophic. Derives its en- 
ergy by the oxidation of thiosulfate to 
sulfate and sulfur ; also oxidizes sulfur to 
sulfate. 

Aerobic. 

Source : Sea water, river water, mud, 
sewage, and soil. 

Habitat : Presumably widely distrib- 
uted. 

2. Thiobacillus thiooxidans Waksman 
and Joffe. (Jour. Bact., 7, 1922, 239; 
Sulfomonas thiooxidans Waksman, Jour. 
Bact., 7, 1922, 616; Thiobacterium thiooxy- 
dans Lehmann and Neumann, Bakt. Diag., 
7 Aufl., 2, 1927, 517.) From Greek 
theion, sulfur and M.L. to oxidize. 

Short rods: 0.5 by 1.0 micron with 
rounded ends. Occur singly, in pairs, 
or in chains . Motile by means of a single 



polar flagellum. Gram-negative (Star- 
key, Soil Sci., 55, 1935,210). 

Thiosulfate agar: Scant growth. 
Nearly transparent colonies. 

Sulfur medium (liquid) : Uniform tur- 
bidity. No sediment or surface growth. 
Medium becomes very acid (below pH 
1.0). 

Thiosulfate medium (liquid) : Uniform 
turbidity. Medium becomes acid and 
sulfur is precipitated. 

Nitrogen sources: Utilizes ammonia 
nitrogen but not nitrate nitrogen which 
is toxic. Asparagin, urea and peptone 
not utilized. 

Temperature relations : Optimum 28° 
to 30°C. Slow growth at 18° and 37°C. 
Death occurs at 55° to 60°C. 

Optimum reaction: pH 2.0-3.5. 
(Limiting reactions, pH 6.0 to less than 
pH 0.5.) 

Strictly autotrophic, deriving its 
energy from the oxidation of elementary 
sulfur and thiosulfate, oxidizing these to 
sulfuric acid. It utilizes the CO2 of the 
atmosphere as a source of carbon. 

Strictly aerobic. 

Distinctive characters : This species 
produces more acid, from oxidation of 
sulfur, and continues to live in a more 
acid medium, than any other living or- 
ganism yet reported, the hydrogen-ion 
concentration of the medium increasing to 
a pH 0.6 and less. 

Source : Isolated from composts of soil, 
sulfur, and rock phosphate and soils con- 
taining incompletely oxidized sulfur 
compounds. 

Habitat: Soil. 



80 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



3. Thiobacillus novellus Starkey. 
(Jour. Bact., 28, 1934, 365; Jour. Gen. 
Physiol., 18, 1935, 325; Soil Sci., 89, 
1935, 207, 210.) From Latin novellus, 
new . 

Short rods or ellipsoidal cells : 0.4 to 
0.8 by 0.6 to 1.8 microns. Non-motile. 
Gram -negati ve . 

Gelatin stab : Mucoid growth at point 
of inoculation. Sub-surface growth 
meager. Slow liquefaction. 

Agar plate: Growth slow, colorless, 
moist, raised, circular, 1 mm in diameter. 
Deep colonies tiny, lens-shaped. 

Thiosulfate agar plate: Growth slow, 
becoming white from precipitated sulfur. 
Surface colonies small, circular, moist. 
Crystals of CaS04 appear throughout the 
agar. 

Agar slant: Growth fairly abundant, 
soft, somewhat ropy, raised, shining, 
moderately spreading; whitish in re- 
flected light, brownish opalescence in 
transmitted light. 

Thiosulfate agar slant : Growth very 
thin, practically colorless. No sub-sur- 
face growth. Sulfur usually precipitated 
as white frosty film on the surface. 

Agar stab : White to cream-colored 
growth confined close to point of inocula- 
tion. Penetrates to bottom of tube. 

Thiosulfate agar stab : No appreciable 
surface growth. 

Broth: Slightly turbid. Gelatinous 
pellicle. Forms long streamer-like 
network extending from surface to the 
bottom. Some sediment. 

Thiosulfate solution medium : Uniform 
turbidity. No pellicle. Whitish sedi- 
ment with thin incomplete membrane 
on the bottom of the flask. Reaction 
acid in a few days, changes pH 7.8 to 5.8 
with decomposition of a small quantity 
of thiosulfate. 

Sulfur solution medium of slightly 
alkaline reaction: No growth. 

Potato slant: Growth limited, cream- 
colored, moist, shining, slightly brown. 

Litmus milk: Slow development of 
slight alkalinity. 

Facultative autotrophic. 



Optimum reaction : Close to neutrality 
(limiting reactions pH 5.0 to 9.0). 

Aerobic. 

Distinctive characters : Oxidizes thio- 
sulfate to sulfate and sulfuric acid. Does 
not oxidize free sulfur. 

Source: Isolated from soils. 

Habitat: Soils. 

4. Thiobacillus coproliticus Lipman 
and McLces. (Soil Sci., 50, 1940, 432.) 
Latinized form of the English word copro- 
lite, fossil dung. 

Long thin rods: 0.1 to 0.2 by 6 to 
8 (may measure 3 to 40) microns. 
Straight, S -shaped, and curved cells. 
Motile by means of a single polar flagel- 
lum. 

Peptone soil extract agar : Slight 
growth. 

Nutrient solution : Little or no growth. 

Thiosulfate agar: Slow development. 
Produces small watery colonies raised 
above the agar surface. Colonies have 
been noted which were white from pre- 
cipitated sulfur. 

Thiosulfate solution: Thiosulfate is 
oxidized. Little or no turbidity. No 
pellicle. No sediment. Change in re- 
action from pH 7.6 to 6.1. 

Sulfur medium: Sulfur is oxidized. 
No turbidity. 

Facultative autotrophic. 

Aerobic. 

Distinctive characters: Develops in 
inorganic media and oxidizes thiosulfate 
and sulfur to sulfate. Media with 
slightly alkaline reactions most favorable 
for growth. 

Source : Coprolite rock material from 
Triassic period (Arizona). 

Habitat : Unknown. 

5. Thiobacillus denitrificans Beijer- 
inck. (Cent. f. Bakt., II Abt., 11, 1904, 
597; Sulfomonas denitrificans Orla-Jen- 
sen. Cent, f . Bakt., II Abt., 22, 1909, 314.) 
From Latin, de, from; and M.L. nitrifico, 
to nitrify. 

Short rods, 0.5 by 1 to 3.0 microns long. 
Motile by means of a single polar flagel- 



FAMILY NITROBACTERIACEAE 



81 



lum (Tjulpanova-Mossevitch, Arch. d. 
Sci. Biol., U.S.S.R., 30, 1930, 203). 

Inorganic liquid medium : Growth witli 
production of gas, predominantly nitro- 
gen. 

Thiosulfate agar medium : Colonies 
thin, clear, or weakly opalescent. 

Optimum reaction : Neutral or slightly 
alkaline. 

Autotrophic, utilizing carbon from CO2, 
carbonates and bicarbonates. Consid- 
ered to be strictly autotrophic by Lieske 
(Ber. d. deutsch. botan. Gesell., 30, 1912, 
12.) and facultative by Tjulpanova- 
Mossevitch {loc.cit.). Beijerinck stated 
(Kon. Akad. v. Wetenschappen Amster- 
dam, 4£, 1920, 899) that whereas the 
organism developed initially in an in- 
organic medium, it lost the autotrophic 
habit by cultivation in an organic me- 
dium. 

Facultative anaerobic or even micro- 
aerophilic. Can live in the absence of 
free O2 in the presence of nitrate. 

Distinctive characters: Oxidizes thio- 
sulfate to sulfate under anaerobic con- 
ditions using nitrate a.s the hydrogen 
acceptor which is reduced to N2. Also 
oxidizes sulfide, elemental sulfur, and 
dithionatel 

Habitat : Canal and river water, salt 
water, soil, peat, composts and mud. 

Appendix: The following species have 
been placed in Thiobucilhis or are re- 
garded as belonging to the genus : 

Thiobacillus concretivorus Parker. 
(Austral. Jour. Exper. Biol, and Med. 
Sci., 23, 1945, SI.) From corroded 
concrete sewers. Similar to or identical 



witli Thiobacillus thiooxidans Waksman 
and JofTe. 

Thiobacillus crenalus Emoto. (Proc. 
Imp. Acad. Tokyo, 5, 1929, 149.) Iso- 
lated from mud of hot springs in Japan. 
See description. Manual, 5th ed., 1939, 
84. Almost identical with Thiobacillus 
thiooxidans Waksman and Joffe. 

Thiobacillus lobatus Emoto {loc. 
cit., p. 148). Source and relationships as 
above. See description, Manual, 5th ed., 
1939, 83. 

Thiobacillus thermilanus Emoto. 
(Bot. Mag. Tokyo, 42, 1928, 422.) 
Source and relationships as above. See 
description. Manual, 5th ed., 1939, 83. 

Thiobacillus trautweinii Bergey et al. 
See Flavobacterium appendix. 

Thiobacillus umbonatus Emoto {loc. 
oil., p. 150). Source and relationships as 
above. See description, Manual, 5th 
ed., 1939,84. 

Thiobaclerium beijerinckii Issat- 
chenko and Salimowskaja. (Zur Mor- 
phologie u. Physiol, der Thionsaure- 
bakterien (Russian with German 
abstract), Izyiestia Gosud. Gidrobiol. 
Inst., No. 21, 1928, 61.) From salt seas in 
Russia. Similar to or identical with 
Thiobacillus thioparus Beijerinck. 

Thiobaclerium beijerinckii var. jacob- 
senii Issatschenko and Salimowskaja {loc. 
cil.). Variety of previously mentioned 
species. 

Thiobaclerium nalhansonii Issat- 
chenko and Salimowskaja {loc. cil.). 
From salt seas in Russia. Similar to or 
identical with Thiobacillus thioparus 
Beijerinck. 



82 MANUAL OF DETERMINATIVE BACTERIOLOGY 

FAMILY II. PSEUDOMONADACEAE WINSLOW ET AL. 
(Jour. Bact., 2, 1917, 555.) 

Cells without endospores, elongate rods, straight or more or less spirally curved. 
One genus (Mycoplana) has branched cells. Usually motile by polar flagella which 
are either single or in small or large tufts. A few species are non-motile. Gram-nega- 
tive (a few doubtful Gram-positive tests are recorded in Pseudomonas). Grow 
well and fairly rapidly on the surface of ordinary culture media excepting Methano- 
monas and some vibrios that attack cellulose. They are preferably aerobic, only 
certain vibrios including Desulfovibrio being anaerobic. Either water or soil forms, 
or plant or animal pathogens. 

Key to the tribes of family Pseudomonadaceae. 

1. Straight rods. 

Tribe I. Pseudomonadeae, p. 82. 

2. Cells more or less spirally curved. 

Tribe II. Spirillcae, p. 192. 

TRIBE I. PSEUDOMONADEAE KLUYVER AND VAN NIEL. 

(Cent. f. Bakt., II Abt., 94, 1936, 397.) 

This tribe includes all of the straight and branching rods of the family. 
Key to the genera of tribe Pseudomonadeae. 
I. Soil and water bacteria. Few animal and many plant pathogens. Usually 
produce a water-soluble pigment which diffuses through the medium as a 
bluish-green or yellowish-green pigment. 

Genus I. Pseudomonas, p. 82. 
II. Cells usually monotrichous with yellow non-water-soluble pigment. Mostly 
plant pathogens causing necrosis. 

Genus II. Xanthomonas, p. 150. 

III. Soil bacteria which oxidize methane. 

Genus III. Methanomonas, p. 179. 

IV. Bacteria which oxidize alcohol to acetic acid. 

Genus IV. Acetobacier, p. 179. 
V. Soil and water bacteria known to attack protamines. 

Genus V. Protaminobacter, p. 189. 
VI. Soil bacteria with branching cells. Capable of using aromatic compounds, 
as phenol, etc., as a source of energy. 

Genus VI. Mycoplana, p. 191. 

Genus I. Pseudomonas Migula* 

(Migula Arb. bakt. Inst. Karlsruhe, 1, 1894, 237; Bacterium Ehrenberg emend. 
Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 1, 1872, 167; BactriUum Fischer, Jahrb. f. 
wissensch. Bot., 27, 1895, 139; Bactrinium Fischer, ibid., 41 ; Arthrobactrinium Fischer, 
ibid., 139; Arthrobactr ilium Fischer, ibid., 139; Bactrinius Kendall, Public Health, 
28, 1902, 484; Bacirillius Kendall, ibid.; Bacterium Ehrenberg emend. Smith, Bacteria 

* Revised for the 5th ed. of the Manual by Prof. D. H. Bergey, Philadelphia, Penn- 
sylvania, 1937. Further revision for the 6th ed. by Prof. R. S. Breed, New York State 
Experiment Station, Geneva, New York, with incorporation of the plant pathogenic 
species by Prof. Walter H. Burkholder, Cornell University, Ithaca, New York, 
April, 1943. 



FAMILY PSEUDOMOXADACEAE 83 

in Relation to Plant Disease, 1, 1905, 171; Dcnilromonas Orla-Jensen, Cent. f. Bakt., 
II Abt., 22, 1909, 314; Liquidomonas Orla-Jensen, ibid., 332; Lamprella Enderlein, 
Sitzber. Gesell. naturf. Freunde, Berlin, 1917, 317; Fluoromonas Orla-Jensen, Jour. 
Bact.. 6, 1921, 271.) 

Cells nionotrichous, lophotrichous or non-motile. If pigments are produced, they 
are of greenish hue, fluorescent, and water-soluble.* Gram-negative except Nos. 
88, 122 and 128. Frequently ferment glucose, sometimes with the formation of 
visible gas. Inactive in the fermentation of lactose. Nitrates are frequently re- 
duced either to nitrites or ammonia, or to free nitrogen. Some species split fat and 
attack hydrocarbons. Soil, water, and plant pathogens; very few animal pathogens. 
Certain salt water species (Nos. 58-64) some of which live in heavy brine are tempo- 
rarily retained in this genus although they produce non-water-soluble pigments or 
phosphorescence. From Gr. pseudes, false; monas, a unit; M. L. monad. 

The type species is Pseudomonas aeruginosa (Schroeter) Migula. 

Key to the species of genus Pseudomonas. 

I. Soil and fresh water forms with a few that are pathogenic on cold or warm 
blooded animals. 
1. Green fluorescent pigment produced, 
a. Gelatin liquefied. 

b. Polar flagellate. 

c. Grow readily at 37°C. Usually bluish-green. 

1. Pseudomonas aeruginosa. 

2. Pseudomonas jaegeri. 
cc. Grow poorly or not at all at 37°C. 

d. Milk not coagulated becoming alkaline. 

e. Soil and water organisms. Not known to 
digest cellulose. 

3. Pseudomonas fluorescens. 

4. Pseudomonas viscosa. 

5. Pseudomonas fairmountensis. 

6. Pseudomonas ureae. 

7. Pseudomonas pavonacea. 
ee. Soil forms that attack cellulose. 

8. Pseudomonas effusa. 
eee. Pathogenic for lizards. 

9. Pseudomonas reptilivorous. 

dd. ]Milk unchanged becoming blue in association with 
lactic acid bacteria. 

10. Pseudomonas syncyanea. 
ddd. Milk coagulated. 

11. Pseudomonas schuylkilliensis. 

12. Pseudomonas chlororaphis. 

13. Pseudomonas myxogenes. 

14. Pseudomonas septica. 
dddd. Soil form. Action on milk not recorded. 

15. Pseudomonas boreopolis. 

* See Tobie, Jour. Boct., 49, 1945, 459 for a discussion of the nature of 
these pigments. 



84 MANUAL OF DETERMINATIVE BACTERIOLOGY 

bb. Non-motile. 

c. CJrows readily at 37°C. 

16. Pscudomonas smaragdina. 
cc. Grows poorly or not at all at 37°C. 

17. Psevdomonas chlorina. 
aa. Gelatin not liquefied. 

b. Polar flagellate. 

c. Grow readily at 37°C. Usually bluish-green. 

18. Pseudomonas oleovorans. 

19. Pseudomonas incognita. 

20. Pseudomonas convexa. 

21. Pseudomonas mildenbergii. 
cc. Grow poorly or not at all at 37°C. 

d. Milk not coagulated. 

22. Pseudomonas putida. 

23. Pseudomonas scissa. 

24. Pseudomonas ovalis. 

25. Pseudomonas striata. 

26. Pseudomonas denitrificans. 
dd. Milk coagulated. 

27. Pseudomonas solaniolens. 
bb. Non -motile. 

c. Grows poorly or not at all at 37° C. 
d. Milk not coagulated. 

28. Pseudomonas eisenbergii. 
2. Green fluorescent pigment not produced or not reported. 

a. Gelatin liquefied, 
b. Polar flagellate. 

c. Grow poorly or not at all at 37°C. No visible gas from 
sugars. 
d. Rapid reduction litmus. Putrid odor. 

29. Pseudomonas puirefaciens . 
dd. Slow reduction litmus. Alkaline. 

30. Pseudomonas mephitica. 

31. Pseudomonas geniculala. 
ddd. Acid coagulated. 

32. Pseudomonas fragi. 

cc. Acid and visible gas from glucose. Optimum tempera- 
ture variable, 
d. Litmus milk reduced and alkaline. 

33. Pseudomonas nehulosa. 
dd. Litmus milk acid coagulated. 

34. Pseudomonas roadunata. 

35. Pseudomonas nmUistriata. 

36. Pseudomonas punctata. 

37. Pseudomonas hydrophila. 

38. Pseudomonas ichthyosmia. 
aa. Gelatin not liquefied. 

b. Polar flagellate. 
c. Grow at 37 °C. 



FAMILY PSEUDOMONADACEAE 85 

39. Pseiidomonas ambigua. 

40. Pseudomonas sinuosa. 

41. Pseudomonas cruciviae. 
cc. Grow poorly or not at all at 37°C. 

d. Action on h3'drocarbons and cellulose unknown. 

42. Psexidomonas rngosa. 
dd. Utilize hydrocarbons. 

43. Pseudomonas desmolyiicum . 

44. Pseudomonas rathonis. 

45. Pseudomonas dacunhae. 

46. Pseudomonas arvilla. 

47. Pseudomonas salopium. 
ddd. Utilize cellulose. 

48. Pseudomonas minuscula. 

49. Pseudomonas tralucida. 

50. Pseudomonas mira. 

aaa. Action on gelatin not recorded. Produces alcoholic fermentation of 
glucose. 

51. Pseudomonas lindneri. 
II. Sea water to brine species. Some species phosphorescent. 

a. Gelatin liquefied. 

b. Polar flagellate. 

c. From sea water. Not deeply pigmented. 

d. Nitrites not produced from nitrates. 

52. Pseudomonas membranoformis . 

53. Pseudomonas marinoglutinosa. 

dd. Nitrites produced from nitrates so far as known, 
e. Digest agar. 

54. Pseudomonas gelatica. 

ee. Deposit calcium carbonate in sea water gelatin 
and agar media in old cultures. 

55. Pseudomonas calcis. 

56. Pseudomonas calciprecipitans . 
eee. Causes skin lesions in marine fish. 

57. Pseudomonas ichlhyodermis . 

cc. Produce highly colored pigments in media containing salt 
or in heavy brines, 
d. Blackens salted butter. 

58. Pseudomonas nigrificans. 

dd. Causes purple discoloration of salted beans. 

59. Pseudomonas beijerinckii. 

ddd. Reddens heavy brines (more than 18 per cent salt). 

60. Pseudomonas salinaria. 

61. Pseudomonas cutirubra. 

ccc. Phosphorescent bacteria from decaying fish and crusta- 
ceans, and phosphorescent organs of sea animals, 
d. Gelatin liquefied. 

62. Pseudomonas harvei/i. 
dd. Gelatin not liquefied. 

63. Pseudomonas phosphorescens . 

64. Pseudomonas pierantonii. 



86 MANUAL OF DETERMINATIVE BACTERIOLOGY 

III. Plant pathogens, causing leaf spot, leaf stripe and similar diseases. 
1. Green fluorescent pigment produced, 
a. Gelatin liquefied. 

b. Acid from sucrose. 

c. Nitrites produced from nitrates. 

65. Pseudomonas martyniae. 

66. Pseudomonas striafaciens. 

67. Pseudomonas tomato. 
cc. Nitrites not produced from nitrates. 

d. Growth in 5 per cent salt. 

68. Pseudomonas aceris. 

69. Pseudomonas angulala. 

70. Pseudomonas aptata. 

71. Pseudomonas primulae. 

72. Pseudomonas viridilivida. 
dd. No growth in 5 per cent salt. 

e. Beef peptone agar turns brown. 

73. Pseudomonas delphinii. 
ee. Beef peptone agar uncolored. 

74. Pseudomonas herhcridis. 

75. Pseudomonas coronafaciens . 

75a. Pseudomonas coronafaciens var. atro- 
pupurea. 

76. Pseudomonas lachrymans. 

77. Pseudomonas maculicola. 

78. Pseudomonas marginata. 

79. Psexidomonas medicaginis. 
79a. Pseudomonas phaseolicola. 

80. Pseudomonas pisi. 

81. Pseudomonas syringae. 
d(i(l. Growth in salt solutions not recorded. 

82. Pseudomonas airofaciens. 

83. Pseudomonas cumini. 

84. Pseudomonas desaiana. 

85. Pseudomonas erodii. 

86. Pseudomonas apii. 

87. Pseudomonas matthiolae. 

88. Pseudomonas mors-pr unormn . 

89. Pseudomonas rimaefaciens. 

90. Pseudomonas papulans. 

91. Pseudomo7ias pseudozoogloeae. 

92. Pseudomonas tabaci. 
ccc. Nitrite production not reported. 

93. Pseudomonas lapsa. 
bb. No acid from sucrose. 

c. Nitrites produced from nitrates. 

94. Pseudomonas bowlesiae. 

95. Pseudomonas intyhi. 

96. Pseudomonas marginalis. 

97. Pseudomonas setariae. 



FAMILY PSEUDOMONADACEAE 87 

cc. Nitrites not produced from nitrates, 
d. Lipolytic. 

98. Pseudomonas polycolor. 
dd. Not lipolytic. 

99. Pseudomo7ias viridiflava. 

99a. Pseiidomo?ias viridiflava var. concen- 
trica. 
ddd. Lipolytic action not reported. 

100. Pseudomonas ananas. 
lOL Pseudomonas ligustri. 

102. Pseudomonas sesami. 

103. Pseudomonas tolaasii. 
bbb. Acid from sucrose not reported. 

c. Nitrites produced from nitrates, 
d. Motile. 

104. Pseudomonas xanthochlora. 
dd. Non-motile. 

105. Pseudomonas rhizoclonia. 
cc. Nitrites not produced from nitrates. 

106. Pseudomonas barkeri. 

107. Pseudomonas gladioli. 
lOS. Pseudomonas mellea. 

ccc. Nitrite production not reported. 

109. Pseudomonas betlis. 

110. Pseudomonas panacis. 
a. Gelatin not liquefied. 

b. Acid from sucrose. 

c. Nitrites produced from nitrates. 

111. Pseitdoinonas alcuritidis. 
cc. Nitrites not produced from nitrates. 

112. Pseudomonas glycinea. 

112a. Pseudomonas glycinea var. japonica. 

113. Pseudomonas savastanoi. 

113a. Pseudomonas savastanoi var. fra.zini. 

114. Pseudomonas lonclliana. 
bb. No acid from sucrose. 

c. Nitrites not produced from nitrates. 

115. Pseudomonas calcndulae. 

116. Pseudomonas cichorii. 

117. Pseudomonas cissicola. 

118. Pseudomonas nectaropliila. 

119. Pseudomonas vihurni. 
bbb. Acid from sucrose not reported. 

c. Nitrites not produced from nitrates. 

120. Pseudomonas mori. 

121. Pseudomonas siizolobii. 

122. Pseudomonas viciae. 

2. Green fluorescent pigment not produced or not reported, 
a. Gelatin liquefied. 

b. Acid from sucrose. 



MANUAL OF DETERMINATIVE BACTERIOLOGY 

c. Nitrites produced from nitrates. 

d. Beef-peptone agar turns dark brown. 

123. Pseudomonas alliicola. 

124. Pseudomonas gardeniae. 

dd. Beef-peptone agar remains uncolored or liglit dis- 
coloration after several weeks, 
e. Colonies tan to brown. 

125. Pseudomonas caryophylli. 

126. PseudomoJias solanacearum. 

126a. Pseudomonas solanacearum var. asia- 
tica. 
ee. Colonies white or colorless. 

127. Pseudomonas castaneae. 

128. Pseudomonas seminum. 
cc. Nitrites not produced from nitrates. 

129. Psendomonas passifloriae. 
bb. No acid from sucrose. 

130. Pseudomonas fabae. 
bbb. Acid from sucrose not reported. 

c. Nitrites not produced from nitrates. 

131. Pseudomonas astragali. 

132. Pseudomonas colurnae. 

133. Pseudomonas maublancii. 

134. Pseudomonas polygoni. 
cc. Nitrate production not reported. 

135. Pseudomonas iridicola. 

136. Pseudomonas levistici. 

137. Pseudomonas radiciperda. 
aa. Gelatin not liquefied. 

b. Acid from sucrose. 

c. Nitrites not produced from nitrates. 

138. Pseudomonas melaphthora. 
cc. Gas from nitrates. 

139. Pseudomonas helianthi. 
bb. No acid from sucrose. /- 

c. Nitrites produced from nitrates. 

140. Pseudomonas alboprecipitans . 

141. Pseudomonas peiasilis. 

142. Pseudomonas lignicola. 
cc. Nitrites not produced from nitrates. 

143. Pseudomonas andropogoni. 

144. Pseudomonas woodsii. 
bbb. Acid from sucrose not reported. 

c. Nitrites produced from nitrates. 

145. Pseudomonas panici-miliacei. 

146. Pseudomonas saliciperda. 
cc. Nitrites not produced from nitrates. 

147. Pseudomonas eriobotryae. 
aaa. Gelatin liquefaction not reported. 

b. Nitrites not produced from nitrates. 

148. Pseudomonas wieringae. 



FAMILY PSEUDOMONADACEAE 



89 



1. Pseudomonas aeruginosa (Schroe- 
ter) Aligula. (Bacterium aeruginosum 
Schroeter, in Cohn, Beitrage z. Biologie, 
1, Heft 2, 1872, 126; Bacillus pyocyaneus 
Gessard, Compt. rend. Acad. Sci., Paris, 
94, 1882, 536; Micrococcus pyocyaneus 
Zopf, Spaltpilze, 2 Aufl., 1884, 83; Bacil- 
lua acruginosus Trevisan, Atti Accad. 
Fis.-Med. Stat., Milano, Ser. 4, 3, 1885, 
11; Bacillus fluorescens Crookshank, 
Man. of Bact., 3rd ed., 1890, 247; not 
Bacillus fluorescens Bergey et al., Man- 
ual, 1st ed., 1923, 287; Pseudomonas 
pyocyanea Migula, in Engler and Prantl, 
Die natiirl. Pflanzenfam, 1, la, 1895, 29; 
Bacterium pyocyaneum Lehmann and 
Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 
267; Migula, Syst. Bakt., 2, 1900, 884.) 
From Latin, full of copper rust, or ver- 
digris; green. 

Rods: 0.5 to 0.6 by 1.5 microns, oc- 
curring singly, in pairs and short chains. 
Motile, possessing one to three polar 
flagella. Monotrichous (Reid, Xaghski, 
Farrell and Haley, Penn. Agr. Exp. Sta., 
Bull. 422, 1942, 6). Gram-negative. 

Gelatin colonies : Yellowish or greenish- 
yellow, fringed, irregular, skein-like, 
granular, rapidly liquefying. 

Gelatin stab : Rapid liquefaction. 
The fluid assumes a yellowish-green or 
bluish-green color. 

Agar colonies: Large, spreading, gray- 
ish mth dark center and translucent 
edge, irregular. Medium greenish. 

Agar slant: Abundant, thin, white, 
glistening, the medium turning green to 
dark brown or black, fluorescent. 

Broth: Marked turbidity with thick 
pellicle and heavy sediment. Medium 
yellowish-green to blue, mth fluores- 
cence, later brownish. Produces pyo- 
cyanin, fluorescein and pjTorubrin (Am. 
Jour. Hyg., 5, 1925, 707). 

Litmus milk: A soft coagulum is 
formed, with rapid peptonization and 
reduction of litmus. Reaction alkaline. 

Potato: Luxuriant, dirty-brown, the 
medium becoming dark green. 

Indole usually not formed (Sandiford, 
Jour. Path, and Bact., U, 1937, 567). 



Nitrates are reduced to nitrites and 
nitrogen. 

Glucose, fructose, galactose, arab- 
inose, maltose, lactose, sucrose, dextrin, 
inulin, glycerol, mannitol and dulcitol 
are not attacked. Acid from glucose 
(Sandiford, loc. cit.). 

Blood serum : Liquefied. Yellow 
liquid, greenish on surface. 

Blood hemolyzed. 

Cultures have marked odor of tri- 
methylamine. 

Aerobic, facultative. 

Optimum temperature 37°C. 

Pathogenic for rabbits, guinea pigs, 
rats and mice. 

Common name : Blue pus organism. 

Source: Pus from wounds. Regarded 
as identical with one of the plant patho- 
gens {Pseudomonas polycolor) by Elrod 
and Braun (Jour. Bact., U, 1942, 633). 

Habitat: Cause of various human and 
animal lesions. Found in polluted 
water and sewage. 

2. Pseudomonas jaegeri Migula. 
{Bacillus proteus fluorescens H. Jaeger, 
Ztschr. f. Hyg., 12, 1892, 593; Migula, 
Syst. d. Bakt., 2, 1900, 885; Bacillus 
proteus-fluorescens Holland, Jour. Bact., 
5, 1920, 220; Proteus fluorescens Holland, 
ibid., 224; Pseudomonas prolea-fluorescens 
Holland, ibid., 224.) Named for H. 
.Jaeger who first described the species. 

Short, thick rods, with rounded ends, 
occurring singly and in pairs. Motile 
with a tuft of polar flagella which may be 
pushed to one side where cells remain 
in a chain. Gram-negative. 

Gelatin colonies: Small, transparent, 
becoming proteus-like. 

Gelatin stab: Marked surface growth. 

Saccate to infundibuliform liquefaction. 
Liquefied portion green fluorescent. 

Agar slant : Thick, yellowish-white 
layer, the medium becoming greenish- 
fluorescent. At times gas is formed. 

Broth : Turbid, with greenish-gray pel- 
licle and sediment. 

Litmus milk : Not coagulated. 



90 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Potato: Thick, pale yellow becoming 
dark brown layer, slimy. The medium 
becomes bluish-gray. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 37°C. 

Pathogenic for mice. 

Source: Regarded by Jaeger as the 
cause of Weil's disease (infectious jaun- 
dice) as it was found repeatedly in 
patients suffering from this disease. 
See Leptospira icterohaemorrhagiae. 

Habitat: Water. 

3. Pseudomonas fluorescens Migula. 
(Bacillus fluorescens liquefaciens Flugge, 
Die Mikroorganismen, 18S6, 289; Migula, 
in Engler and Prantl, Die natiirl. Pflan- 
zenfamilien, 1, la, 1895, 29; Bacterium 
fluorescens Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., .2, 1896, 272.) From Latin, 
fluor, flowing; M.L. fluoresco, to fluoresce. 

Rods: 0.3 to 0.5 by 1.0 to 1.8 microns, 
occurring singly and in pairs. Motile, 
possessing a polar flagellum. Gram- 
negative. 

Gelatin colonies: Circular, with green- 
ish center, lobular, liquefying quickly. 

Gelatin stab : Infundibuliform lique- 
faction, with whitish to reddish-gray 
sediment. 

Agar slant : Abundant, reddish layer, 
becoming reddish-gray. The medium 
shows greenish to olive-brown coloration. 

Broth: Turbid, flocculent, with yellow- 
ish-green pellicle and grayish sediment. 

Litmus milk: No coagulation; becom- 
ing alkaline. 

Potato: Thick, grayish-yellow, spread- 
ing, becoming light sepia-brown in color. 

Indole is not formed. 

Nitrates reduced to nitrites and am- 
monia. 

Acid from glucose. 

Blood serum liquefied. 

Aerobic. 

Optimum temperature 20° to 25°C. 

Not pathogenic. 

Source: Water, sewage, feces. 

Habitat: Soil and water. 



4. Pseudomonas viscosa (Frankland 
and Frankland) Migula. (Bacillus vis- 
cosKs G. and P. Frankland, Ztschr. f. 
Hyg., 6, 1889, 391 ; Migula, Syst. d. Bakt., 
2, 1900, 900.) From M. L. viscidus, 
sticky, viscid. 

Small rods: 0.5 by 1.5 to 2.0 microns, 
occurring singly. Motile and presum- 
ably polar flagellate. Gram-negative. 

Gelatin colonies: Grayish, granular, 
with fimbriate margin. Medium assumes 
a green fluorescent color around each 
colony. 

Gelatin stab: Infundibuliform lique- 
faction. Liquefied portion green fluores- 
cent with greenish-white pellicle. 

Agar slant: Thin, greenish-white, the 
medium becoming greenish. 

Broth: Turbid, with greenish pellicle. 

Litmus milk: Not coagulated. 

Potato : Moist, chocolate-brown, viscid. 

Indole not formed. 

Nitrites not produced from nitrates. 
Destroys nitrate with the production of 
ammonia. 

Aerobic, facultative. 

Distinctive characters : Resembles 
Pseudomonas fluorescens except that 
growth on agar, gelatin and potato is 
viscid. 

Optimum temperature 20°C. 

Source: Unfiltered water from Kent, 
England. Common. 

Habitat: Water. 

5. Pseudomonas fairmountensis 

(Wright) Chester. (Bacillus fairmoun- 
tensis Wright, Memoirs Nat. Acad. 
Sci., 7, 1895, 458; Chester, Man. 
Determ. Bact., 1901 , ill ; Achrojuobacler 
Jairmountense Bergey et al., Manual, 
1st ed., 1923, 146.) From M. L. of Fair- 
mount Park (Philadelphia.) 

Medium-sized rods, occurring singly, 
in pairs and in chains. Motile, posses- 
sing polar flagella. Gram-negative. 

Gelatin colonies : Circular, white, 
translucent. Dark centers with a green- 
ish shimmer, thinner edges and faint 
radial lines. 

Gelatin stab : Crateriform liquefaction. 



FAMILY PSEUDOMONADACEAE 



91 



Agar slant : Grayish-white, glistening. 
Broth: Turbid. 

Litmus milk : Alkaline, litmus reduced. 
Potato: Raised, granular, spreading, 
viscid. 
Indole is formed. 

Nitrites not produced from nitrates. 
Aerobic, facultative. 
Optimum temperature 20° to 25°C. 
Habitat: Water. 

6. Pseudomonas ureae Bergey et al. 
(Culture No. 3 of Rubentschick, Cent, f . 
Bakt., II Abt., 72, 1927, 101; Bergey et 
al., Manual, 3rd ed., 1930, 173.) From 
Greek, urum, urine; M. L., urea, urea. 

Rods: 0.6 to 0.7 by 1.7 to 2.0 microns, 
occurring singly and in pairs. Motile. 
Gram-positive. 

Gelatin stab : Infundibuliform lique- 
faction. 

Agar colonies: Circular, grayish-white. 

Agar slant : Grayish-white layer be- 
coming greenish-fluorescent. 

Broth: Turbid. 

Litmus milk: Peptonized. 

Potato: Yellowish-brown streak. 

Indole not formed. 

Nitrates reduced with gas formation. 
Ammonia formed. 

Urea attacked. 

Hydrogen sulfide formed. 

Methylene blue reduced. 

Aerobic, facultative. 

Can grow at 0° C. 

Optimum temperature 20°C. 

Habitat : Sewage filter beds. 

This species is included here through 
an oversight. It should have been 
placed in the Appendix to the genu.s 
Pseudomonas as the original description 
is too incomplete to determine its real 
nature. It is reported to be Gram-posi- 
tive and motile; but the number and ar- 
rangement of flagella are not given. If 
it really is Gram-positive, the species is 
probably peritrichous and does not l)e- 
long in Pseudomonas. 

7. Pseudomonas pavonacea Levine 
and Soppeland. (Bui. No. 77, Iowa 



State Agricultural College, 1926, 41.) 
From Latin, pavo, peacock. 

Rods : 0.5 by 4.5 microns, with truncate 
ends, occurring singly and in chains. 
Motile. Gram-negative. 

Gelatin stab: Crateriform liquefaction. 
Medium becoming brown. 

Agar colonies: Circular, raised, becom- 
ing green, amorphous, entire. 

Agar slant : Greenish, smooth, glisten- 
ing, viscid, medium becoming slightly 
brown. 

Broth: Turbid, with viscid sediment. 
Medium turned dark brown. 

Litmus milk: Slightly alkaline. Lit- 
mus reduced. Peptonized after 10 days. 

Potato: No growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Blood serum liquefied in 5 days. 

No acid or gas from carbohydrate 
media. 

Aerobic, facultative. 

Optimum temperature 22°C. 

Source: Isolated from activated 
sludge. 

8. Pseudomonas effusa Kellerman et 
al. (Kellerman, McBeth, Scales and 
Smith, Cent. f. Bakt., II Abt., 39, 1913, 
515; also Soil Science, 1, 1916, 472; Cel- 
Udomonas effusa Bergey et al.. Manual, 
1st ed., 1923, 162; Bacillus effusus ap- 
pears first as a synonym in Bergey at al., 
ibid. ; later used as name of species 5th 
ed., 1939, 610.) From Latin, effusus, ef- 
fuse, spread out. 

Rods: 0.4 by 1.7 microns. Motile 
with one to three polar flagella. Gram- 
negative. 

Gelatin stab: T^iquefaction. 

Agar slant: Luxuriant, glistening, 
moist, creamy growth. Greenish fluo- 
rescence. 

Peptone starch agar slant: Abundant, 
flat, moist rich creamy growth. Medium 
shows greenish fluorescence. 

Broth: Turbid. 

Litmus milk: Alkaline. Coagulation 
and digestion. 



92 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Potato: Rich, creamy spreading 
growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Ammonia is produced. 

Acid from glucose, maltose, starch, 
glycerol and mannitol. No acid from 
lactose or sucrose. 

Cellulose is attacked. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Source: Isolated from soils in Utah. 

Habitat: Soil. 

8a. Psendomonas effusa var. non-lique- 
faciens Kellerman et al. (loc. cit.). A 
non-liquefying variety that acts more 
slowly on litmus milk. 

Source: Soils from Utah. 

9. Pseudomonas reptilivorous Cald- 
well and Ryerson. (Jour. Bact., 39, 1940, 
335.) From Latin, reptile, a reptile and 
voro, to devour, destroy. 

Rods: 0.5 by 1.5 and 2.0 microns, oc- 
curring singly, in pairs and in short chains 
and having rounded ends. Actively 
motile with two to six polar flagella. 
Gram-negative. 

Gelatin colonies : After 24 hours, small, 
circular, smooth, entire. Liquefaction 
with a yellowish-green fluorescence. 

Gelatin stab : Infundibuliform lique- 
faction becoming stratiform. Putrid odor 
present. 

Agar cultures: Circular, smooth, glis- 
tening, slightly raised, butyrous, translu- 
cent, 2 mm in diameter. 

Agar slant: Growth abundant, smooth, 
filiform, glistening, butyrous and translu- 
cent. 

Broth: Turbid with pellicle and sedi- 
ment. Putrid odor. 

Litmus milk: Alkaline, peptonization, 
complete reduction. Disagreeable odor. 

Potato: Growth moderate, spreading, 
glistening, yellowish-gray to creamy. 
Disagreeable odor. Medium becomes 
brownish-gray. 

Indole not formed. 

Nitrates not produced from nitrates. 



Hydrogen sulfide not produced. 

Slightly acid, becoming alkaline in 
glucose. No acid from arabinose, xylose, 
lactose, sucrose, maltose, trehalose, raf- 
finose, mannitol, dulcitol, inositol and 
salicin. 

Starch not hydrolyzed. 

Pathogenic for guinea pigs and rabbits, 
horned lizards, Gila monsters and chuck- 
wallas. 

Temperature relations : Optimum 20° 
to 25°C. Maximum 37°C. 

Distinctive characters : Yellowish- 
green fluorescence present in meat in- 
fusion media. Pathogenic. 

Source : Isolated in a bacterial disease 
of horned lizards and Gila monsters. 

Habitat: Pathogenic for lizards. 

10. Pseudomonas syncyanea (Ehren- 
berg) Migula. {Vibrio syncyaneus Eh- 
renberg, Berichte ii.d. Verh. d. k. Preuss. 
Akad. d. Wissensch. z. Berlin, 5, 1840, 
202 ; Vib7-io cyanogen.es Fuchs, Magazin fiir 
die gesamte Tierheilkunde, 7, 1841, 190; 
Bacillus syncyaneus Schroeter, Krypto- 
gam. Flora von Schlesien, S, 1 , 1886, 157 ; 
Bacillus cyanogenus Zopf, Die Spalt- 
pilze, 3 Auk., 1885, 86; Migula, in Engler 
and Prantl, Die nattirl. Pflanzenfam., 
/, la, 1895, 29; Bacterium syncyaneum 
Lehmann and Neumann, Bakt. Diag., 1 
Aufl., 2, 1896, 275; Pseudomonas cyano- 
genes Holland, Jour. Bact., 5, 1920, 224.) 
From Greek, syn, with; kyaneos, dark 
blue, dark. 

Rods with rounded ends, occurring 
singly, occasionally in chains, 0.7 by 2.0 
to 4.0 microns. Motile with two to four 
polar flagella. Gram-negative. 

Gelatin colonies: Flat, bluish, trans- 
lucent. 

Gelatin stab : Surface growth shiny, 
grayish blue. The medium is colored 
steel-blue with greenish fluorescence. 
Gelatin is liquefied. Some strains do not 
liquefy. 

Agar slant : Grayish-white streak. The 
medium takes on a bluish-gray color with 
slight fluorescence. 



FAMILY PSEUDOMONADACEAE 



93 



Broth: Turbid mth marked fluo- 
rescence. 

Litmus milk: Unchanged. In associa- 
tion with lactic acid bacteria the milk 
takes on a deep blue color. 

Potato: Yellowish-gray, shiny layer, 
becoming bluish-gray. The medium be- 
comes bluish-gray. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: The cause of blue milk. 

11. Pseudomonas schuylkilliensis 

Chester. (Bacillus fluorescens schuyl- 
killiensis Wright, ]\Iemoirs, Xatl. Acad. 
Sci., 7, 1895, 448; Chester, Determinative 
Bact., 1901, 320.) From :\I. T>. of the 
Schuylkill (River). 

Synonyms: Pseudomonas capsulala 
Chester, Man. Dctcrm. Bact., 1901, 322 
(Bacillus fluorescens capsulalus Pottien, 
Ztschr. f. Hyg., 11, 1896, 140); Pseudo- 
monas dennatogenes Fuhrmann, Cent, 
f. Bakt., II .\bt., 17, 1906, 356. 

Short rods, with rounded ends, occur- 
ring singly, in pairs and in chains. Mo- 
tile, possessing a polar flagellum. Gram- 
negative. 

Gelatin colonies: Grayish-white, trans- 
lucent, with brownish center, radiate 
margin, becoming bluish-green. 

Gelatin stab: Slow crateriform lique- 
faction, with blue-green fluorescence. 

Agar slant : Grayish, translucent 
growth. Medium shows greenish fluores- 
cent. 

Broth: Turbid, with slight pellicle and 
blue-green fluorescence. Stringy sedi- 
ment. 

Litmus milk: Coagulated, with slow 
reduction of litmus; peptonized. 

Potato: Brownish, spreading, viscid, 
thick. 

Indole is formed (trace). 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Does not grow at 35° to 36°C. 



Source : Isolated from Schuylkill River 
water. 
Habitat: Water. 

12. Pseudomonas chlororaphis (Guig- 
nard and Sauvageau) Bergey et al. (Ba- 
cillus chlororaphis Guignard and Sauva- 
geau, Compt. rend. Soc. Biol. Paris, 1, 
10 s6r., 1894, 841;. Bergey et al., Manual, 
3d ed., 1930, 183; also see Lasseur and 
Dupaix-Lasseur, Trav. Lab. Microbiol. 
Fac. Pharm. Nancy, Fasc. 9, 1936, 35.) 
From Greek, chloros, greenish yellow; 
r aphis, needle. 

Rods: 0.8 by 1.5 microns, with rounded 
ends, occurring singly and in pairs. Mo- 
tile with polar flagella. Gram-negative. 

Gelatin colonies: Circular, viscid, 
transparent, glistening, lobate margin, 
with fluorescent corona. Dissociates 
readily (Lasseur and Dupaix-Lasseur, 
loc. cit.). 

Gelatin stab: Stratiform liquefaction. 

Broth: Turbid, fluorescent, with crys- 
tals of green, water-soluble chloro- 
raphine. 

Litmus milk : Coagulation. Peptoniza- 
tion. Crystals of chlororaphine form 
in the central part of the culture. 

Potato: Citron-yellow layer. Crystals 
of chlororaphine are formed. 

Nitrates reduced to nitrites. 

Indole not formed. 

Pigment formation: Asparagine, potas- 
sium phosphate, glycerol, sulfate of mag- 
nesium and sulfate of iron are indispensa- 
ble to the formation of crystals of 
chlororaphine. 

Aerobic, facultative. Optimum tem- 
perature 25° to 30°C. 

Pathogenic for laboratory animals. 
Exotoxin formed. 

Habitat: Water. 

13. Pseudomonas myxogenes Fuhr- 
mann. (Cent. f. Bakt., II Abt., 17, 
1907, 356.) From Greek, myxa, mucus; 
gennao, to beget; M. L. slime producing. 

Rods: 0.4 to 0.5 by 1.0 to 1.5 microns, 
occurring singly and in pairs. Motile, 
possessing a bundle of five to seven polar 
flagella. Gram-negative. 



94 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Gelatin colonies: Smooth, soft, flat, 
spreading, entire, yellowish-green. 

Gelatin stab: Growth along stab. 
Liquefaction with yellowish-white sedi- 
ment. 

Agar colonies : Circular, raised, smooth, 
amorphous, entire. 

Agar slant: Yellowish-white, moist, 
glistening, becoming light green-fluores- 
cent. 

Broth: Turbid, with yellowish-white 
sediment. 

Litmus milk: Flocculent precipitation. 
Slow peptonization with yellow serum. 
Alkaline. 

Potato : Dirty yellow to olive, moist, 
glistening, entire. 

Indole is formed. 

Nitrates reduced to nitrites and am- 
monia. No gas formed. 

Aerobic, facultative. 

Optimum temperature 22°C. 

Source: Isolated from beer. 

14. Pseudomonas septica Bergey et al. 
{Bacillus fluorescens septicus Stutzer 
and Wsorow, Cent. f. Bakt., II Abt., 71, 
1927, 113; Bergey et al.. Manual, 3rd ed., 
1930, 169.) From Greek, septikos, putre- 
factive, septic. 

Rods: 0.6 to 0.8 by 0.8 to 2.0 microns, 
occurring singly. Motile with a polar 
flagellum. Gram-negative. 

Gelatin stab: Infundibuliform lique- 
faction. 

Agar colonies: Circular with opalescent 
center and transparent periphery. 

Agar slant : Moderate, undulate margin. 

Broth: Turbid with fragile pellicle, 
greenish in upper poution. 

Litmus milk: Alkaline, coagulated. 

Blood serum not liquefied. 

Acid from glucose. 

Aerobic, facultative. 

Optimum temperature 20^0. 

Habitat: Disease of caterpillars. 

15. Pseudomonas boreopolis Gray and 
Thornton. (Gray and Thornton, Cent. 
f. Bakt., II Abt., 73, 1928, 74.) From 



Greek, boreas, the North wind; polis, 
city; M. L. North City. 

Rods: 0.5 to 1.0 by 2.0 to 3.0 microns, 
occurring singly and in pairs. Motile 
with one to five polar flagella. Gram- 
negative. 

Gelatin colonies : Liquefied. 

Gelatin stab : Liquefied. Medium red- 
dened. 

Agar colonies: Circular or amoeboid, 
white to buff, flat to convex, smooth, 
glistening, translucent border. 

Agar slant: Filiform, whitish, raised, 
smooth, glistening, fluorescent. 

Broth: Turbid. 

Nitrates reduced to nitrites. 

Starch not hydrolyzed. 

Acid produced from glucose. 

Attacks naphthalene. 

Aerobic, facultative. 

Optimum temperature 20° to 25°C. 

Habitat: Soil. 

16. Pseudomonas smaragdina Migula. 
(Bacillus S7naragdimis foetidus Rei- 
man, Inaug. Dissertation, Wiirzburg, 
1887;Migula,Syst.d. Bakt.,.?, 1900, 890.) 
From Greek, smaragdinas, green like the 
smaragdus, the emerald. 

Small rods, occurring singly. Non- 
motile. Gram -negative. 

Gelatin colonies : Small, convex, irregu- 
lar, whitish with greenish shimmer. 

Gelatin stab: Slight surface growth. 
Infundibuliform liquefaction. The 
liquefied medium becomes light emerald 
green in color. 

Agar colonies: Small, brownish-yellow, 
convex. 

Agar slant : Abundant growth with 
greenish fluorescence. 

Broth: Turbid. 

Litmus milk: Not coagulated. 

Potato: Dark brown, becoming choco- 
late brown. 

Indole not formed. 

Nitrates not reduced. 

The cultures give off an odor resembling 
jasmine. 

Aerobic, facultative. 



FAMILY PSEUDOMONADACEAE 



95 



Optimum temperature 37°C. 

Subcutaneous and intravenous inocula- 
tions into rabbits cause death in 36 to 48 
hours. 

Source : Isolated from nasal secretions 
in ozena. 

17. Pseudomonas chlorina (Frankland 
and Frankland) Levine and Soppeland. 
{Bacillus chlorinus G. and P. Frankland, 
Philos. Trans. Roy. Soc. London, 178, 
1887, 274; Bacterium chlorinum Migula, 
Syst. d. Bakt., 2, 1900, 471; Levine and 
Soppeland, Bui. No. 77, Iowa State Agri- 
cultural College, 1926.) From Greek, 
chloros, greenish yellow. 

Piods : 0.5 by 1 .5 micron, occurring singly 
and in short chains. Non-motile. Gram- 
negative. 

Gelatin stab : Crateriform liquefaction 
with green fluorescence. Lemon yellow 
sediment. 

Agar colonies : Circular, raised, smooth, 
amorphous, entire, becoming greenish 
yellow. 

Agar slant : Slightly raised, glistening, 
the medium becoming light greenish 
yellow. 

Broth : Moderate turbidity. Dirtj' 
yellow sediment. No pellicle. 

Litmus milk: Peptonized. Litmus re- 
duced. 

Potato: Scant, olive green growth. 

Indole formed. 

Nitrites produced from nitrates. 

Starch hydrolyzed. 

Blood serum liquefied in .5 days. 

Acid from glucose. 

Aerobic, facultative. 

Optimum temperature 22°C. 

Source : Air. 

18. Pseudomonas oleovorans Lee and 
Chandler. (Jour. Bact., 41, 1941, 378.) 
From M. L. oil destroying. 

Short rods: 0.5 by 0.8 to 1.5 microns, 
occurring singly and in pairs. . Motile. 
Gram-negative. 

Gelatin stab : No liquefaction after 6 
weeks. 



Gelatin colonies: Up to 1 mm. in diam- 
eter, fluorescent ; similar to agar colonies. 

Surface agar colonies : After 24 hours 
1 to 2 mm. in diameter, smooth, convex, 
shin3% opaque, creamy, fluorescent by 
transmitted light. Edge entire in young 
colonies. 

Deep agar colonies: 0.5 by 1.0 to 1.5 
mm., lens-shaped, buff-colored, not flu- 
orescent. 

Agar slant : Growth raised, smooth, 
fluorescent, edge erose. 

Broth : After 24 hours moderate turbid- 
ity with slight yellowish viscid sediment. 
No pellicle or ring. 

Litmus milk: No change. 

Indole not formed. 

Potato : Good growth. 

Nitrites are produced from nitrates. 

Starch is hydrolyzed. 

No acid from glucose, lactose, sucrose, 
galactose, xylose, mannitol, salicin and 
glycerol. 

Equally good growth at 25° and 37°C. 

Aerobic. 

Distinctive character: The fluorescent 
quality of the colonies is not imparted to 
any of the artificial media used. 

Source : Isolated from cutting com- 
pound (oil-water emulsion) circulating 
in a machine shop. The oil in this 
compound may be utilized as a sole source 
of energy. 

Habitat : Probably oil-soaked soils. 
Abundant in cutting compounds. 

19. Pseudomonas incognita Chester. 
{Bacillus Jiuorescens incognilus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 436; 
Chester, Determinative Bacteriology, 
1901, 323.) From Latin, in, not; cogito, 
to think; M. L. unknown. 

Short rods, with rounded ends, occur- 
ring singly, in pairs and in chains. Mo- 
tile, possessing a polar flagellum. Gram- 
negative. 

Gelatin colonies: Thin, transparent, 
slightly granular, becoming greenish. 
Margin undulate. The medium assumes 
a blue-green fluorescence. 



96 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Gelatin stab : No liquefaction. 

Agar slant: Thin, moist, translucent, 
becoming greenish. 

Broth: Turbid, with pellicle, becoming 
greenish. 

Litmus milk: Slightly acid in a month. 
The litmus is slowly reduced. 

Potato: Moist, glistening, spreading, 
brown. 

Indole is formed (trace). 

Nitrites are produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 35°C. 

Habitat: Water. 

20. Pseudomonas convexa Chester. 
{Bacillus fluorescens convexus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 438; 
Chester, Determinative Bacteriology, 
1901, 325.) From Latin, convexus, con- 
vex, arched. 

Short, thick rods, with rounded ends. 
Motile, possessing a polar flagellum. 
Gram -negative. 

Gelatin colonies: Circular, convex, 
glistening, bright greenish, translucent. 
The medium becomes blue-green, flu- 
orescent. 

Gelatin stab: Light green, raised, 
gUstening surface growth. No liquefac- 
tion. 

Agar slant : Moist, translucent, glisten- 
ing, light greenish. The medium as- 
sumes a greenish color. 

Broth: Turbid, becoming greenish. 

Litmus milk: No coagulation; alkaline. 

Potato: Pale brown, spreading. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Habitat: Water. 

21. Pseudomonas mildenbergii Ber- 
gey et al. (Der blaue bacillus, Milden- 
berg. Cent. f. Bakt., II Abt., 56, 1922, 
309; Pseudomonas cyanogena Bergey et 
al.. Manual, 1st ed., 1923, 129; not Bacil- 
lus cyanogenes Flugge, Die Mikroorgan- 



ismen, 1886, 201 ; not Pseudomonas cyano- 
genes Hammer, Dairy Bact., 1928, 70; 
Bergey et al.. Manual, 3rd ed., 1930, 
172.) Named for Mildenberg who first 
isolated this species. 

Rods: 0.3 to 0.5 by 1.0 to 3.5 microns, 
with rounded ends, occurring singly. 
Motile, possessing polar flagella. Gram- 
negative . 

Gelatin colonies: Circular, lobed, 
smooth, glistening, slightly raised, steel- 
blue, entire. 

Gelatin stab: No liquefaction. 

Agar colonies: Small, circular, yellow- 
ish or reddish-yellow, entire, becoming 
lobed, grajash-green, iridescent. The 
medium becomes dirty grayish-green. 

Agar slant: Smooth, spreading, slimy, 
glistening, grayish-green to dark green, 
fluorescent. 

Broth: Turbid green, iridescent to 
opalescent with slimy sediment. 

Litmus milk: Not coagulated, blue 
ring. 

Potato: Slimy, glistening, spreading, 
steel blue. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 25''C. 

Source: Isolated from air. 

2. Pseudomonas putida (Trevisan) 
Migula. {Bacillus Jluorcscens putidus 
Flugge, Die Mila-oorganismen, 2 Aufl., 
1886, 288; Bacillus putidus Trevisan, 
I gen. e le specie d. Batteriacee, 1889, 18; 
Migula, in Engler and Prantl, Die na- 
tiir. Pflanzenfam., 1, la, 1895, 29; Ba- 
cillus fluorescens putridus (sic) Kruse, 
in Flugge, Die Mikroorganismen, 2, 
1896, 292; Bacterium putidum Lehmann 
and Neumann, Bakt. Diag., 1 Aufl., 2, 
1896, 271 ; Pseudomonas putrida (sic) 
Migula, Syst. d. Bakt., 2, 1900, 912.) 
It is not clear which spelling should be 
used. Either is correct. From Latin 
putida or putrida, rotten, stinking. 
Rods, with rounded ends. Motile, 



FAMILY PSEUDOMONADACEAE 



97 



possessing polar flagella. Gram-negative. 

Gelatin colonies : Small, finely granular, 
fluorescent with dark center, surrounded 
by a yellow zone, with pale gray margin. 

Gelatin stab : Dirty-white surface 
growth, becoming greenish, fluorescent. 
No liquefaction. 

Agar colonies : Circular, raised, smooth, 
amorphous, entire, with fluorescent zone 
around the periphery. 

Agar slant : Yellowish-green layer, be- 
coming fluorescent. 

Broth: Turbid, fluorescent. 

Litmus milk: Unchanged. 

Potato: Thin, gray to brownish, slimj^ 
layer. 

Cultures give off odor of trimethyl- 
amine. 

Indole not formed. 

Nitrites are produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 25°C. Will grow 
at 37°C. (Pteid et al., Penn. Agr. Exp. 
Sta., Bull. 422, 1942, 9). 

Distinctive features: Identical with 
Pseudomonas fluorescens Migula accord- 
ing to Lehmann and Neumann {loc. cit.) 
except that it does not liquefy gelatin. 
See Pseudomonas eisenbergii Migula. 

Habitat; Putrefying materials; water. 

23. Pseudomonas scissa (Frankland 
and Frankland) Migula. {Bacillus scis- 
8MS G. and P. Frankland, Ztschr. f. Hyg., 
6, 1889, 398; Migula, Syst. d. Bakt., 2, 
1900, 927.) From Latin, scissus, p.p. of 
scindo, to cut. 

Rods: 0.5 by 0.5 to 1.0 micron, with 
rounded ends, occurring singly, in pairs 
and in chains; on gelatin, coccus-like. 
Motile with presumably polar flagella. 
Gram-negative. 

Gelatin colonies: Small, greenish. 

Gelatin stab : Thin, smooth, ghstening 
surface growth, irregular, serrate margin. 
No liquefaction. The medium becomes 
light green in color. 

Agar slant: Smooth, glistening, lobed. 
The medium assumes a greenish color. 

Broth: Turbid, with whitish sediment. 



Litmus milk : Not coagulated. 

Potato : Glistening, reddish-brown 
growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Distinctive characters : Resembles 
Pseudomonas viscosa Migula. 

Source : Found in water and soils in 
Kent, England. 

Habitat : Water and soil. 

24. Pseudomonas ovalis Chester. 
(Bacillus fluorescens ovalis Ravenel, 
Memoirs Nat. Acad. Sci., 8, 1896, 9; 
Chester, Determinative Bacteriologj^, 
1901, 325; not Bacillus ovalis Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 435.) 
From ovum, egg; M. L. oval. 

Rods: 0.3 to 0.7 by 0.7 to 1.3 microns, 
occurring singly. Motile, possessing a 
single polar flagellum. Gram-negative. 

Gelatin colonies : Irregular, lobate, 
slightly granular. 

Gelatin stab : No liquefaction. 

Agar colonies : Circular, opaque, entire, 
ureenish fluorescence. 

Agar slant: Thick, white, becoming 
greenish^ fluorescent. 

Broth: Turbid, with pellicle. 

Litmus milk : No coagulation ; alkaline. 

Potato: Luxuriant, dirty-brown. 

Indole not formed. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Blood serum not liquefied. 

Acid from glucose. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: Soil. Has been found in in- 
testinal canal. 

25. Pseudomonas striata Chester. 
(Bacillus striatus viridis Ravenel, 
Memoirs Nat. Acad. Sci., 8, 1896, 22; 
Chester, Determinative Bacteriology, 
1901, 325.) From Latin, strio, streak, 
groove. 

Slender rods, of variable lengths, stain- 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



ing irregularly, occurring singly and in 
pairs. Motile, possessing polar flagella. 
Gram -negative. 

Gelatin colonies: Circular, yellowish, 
with filamentous border. 

Gelatin stab: Raised, white surface 
growth. No liquefaction. 

Agar slant: Thin, yellowish-green, 
smooth, glistening. 

Broth: Turbid, becoming slightly 
greenish. 

Litmus milk : No coagulation ; becom- 
ing alkaline; litmus reduced. 

Potato : Moist, glistening, becoming 
chocolate-brown. 

Indole is formed. 

Nitrites are produced from nitrates. 

Aerobic. 

Optimum temperature 25°C. 

Habitat: Soil. 

26. Pseudomonas denitrificans Ber- 
gey et al. {Bacillus denitrificans fluo- 
rescens Christensen, Cent. f. Bakt., II 
Abt., 11, 1903, 190; Bergey etal.. Manual, 
1st ed., 1923, 131.) From Latin, de, 
from, out of; nitrum, soda, nitre; M. L. 
denitrifying. 

Rods: 0.5 to 0.7 by 0.5 to 1.25 microns, 
occurring singly and in pairs in large, 
slimy masses. Motile. Gram-negative. 

Gelatin colonies: Small, circular, con- 
toured, raised, moist, pearly-gray, glis- 
tening. 

Gelatin stab: Whitish, lobed surface 
growth. Yellowish-green growth in stab. 
No liquefaction. 

Agar colonies : Pearly white, circular, 
entire. 

Agar slant: Broad, whitish, contoured, 
moist, entire. 

Broth: Turbid, with thick, wrinkled 
pellicle. 

Litmus milk : Not coagulated. 

Potato: Reddish-gray layer. 

Indole not formed. 

Nitrates reduced with production of 
nitrogen. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: Soil. 



27. Pseudomonas solaniolens Paine. 
(Rept. Int. Conf. Phytopath. and Econ. 
Ent. Holland, 1923, 77; Phytomonas so- 
laniolens Bergey et al.. Manual, 3rd ed., 
1930, 274.) From M. L. Solanum, a 
generic name. 

Small oval rods: Motile with a polar 
flagellum. Gram-negative. 

Produces an iridescence in gelatin. 

Gelatin: No liquefaction. 

Gelatin colonies: Round. Iridescence 
in medium. 

Agar colonies : Pale buff. 

Litmus milk: Curd, with no sign of 
digestion. 

Potato : Pale buff-colored growth, no 
change in medium. 

Nitrites not produced from nitrates. 

Acid but not gas from glucose. No 
acid or gas from lactose, sucrose, manni- 
tol or glycerol. 

Starch: Action feeble. 

Optimum temperature 20° to 30°C. 

Aerobic, facultative. 

Source : Isolated from potato showing 
internal rust spots. 

28. Pseudomonas eisenbergii Migula. 
(Bacillus fluurescens non liquefaciens 
Eisenberg, Bakt. Diag., 3 Aufl., 1891, 
145; Bacillus fluorescens immobilis Kruse, 
in Fliigge, Die Mikroorganismen, 2, 1896, 
294; Migula, Syst. d. Bakt., 2, 1900, 913; 
Bacterium immobile Chester, Man. De- 
term. Bact., 1901, 180; Pseudomonas non- 
liquefaciens Bergey et al., Manual, 1st 
ed., 1923, 132.) Named for Eisenberg, 
the bacteriologist who first reported this 
species. 

Short, slender rods, with rounded ends, 
occurring singly. Non-motile. Kruse 
(loc. cit., p. 293) lists the motile form of 
this organism as Bacillus fluorescens non 
liquefaciens. Gram-negative. 

Gelatin colonies : Fern-like surface 
colonies. Medium around colonies has a 
pearly luster. 

Gelatin stab : Surface growth has fluo- 
rescent shimmer. No liquefaction. 

Agar slant : Greenish layer. 

Broth: Turbid, fluorescent. 



FAMILY PSEUDOMONADACEAE 



99 



Litmus milk : Unchanged. 
Potato : Diffuse, brownish layer. Me- 
dium acquires a grayish-blue color. 
Indole is not formed. 
Nitrites produced from nitrates. 
Acid from glucose. 
Blood serum liquefied. 
Aerobic, facultative. 
Optimum temperature 25°C. 
Not pathogenic. 
Habitat : Water. 

29. Pseudomonas putrefaciens (Derbj^ 
and Hammer) Long and Hammer. 
(Achromobacter putrefaciens Derby and 
Hammer, Iowa Agr. Exp. Sta. Res. Bui. 
145, 1931, 401; Long and Hammer, Jour. 
Bact., 41, 1941, 100.) From Latin, 
piiirida, rotten, stinking; faciens, mak- 
ing. 

Rods: 0.5 to 1.0 by 1.1 to 4.0 microns, 
occurring singly and in pairs. Motile, 
with a single flagellum. Gram-negative. 

Gelatin stab : Rapid, saccate to strati- 
form liquefaction, with reddish-brown 
sediment in the liquefied portion. 

Agar colony : Circular, smooth, glisten- 
ing, slightly raised, somewhat trans- 
parent, with brownish tinge. 

Agar slant : Echinulate, slightly red- 
dish-brown, viscous. 

Broth : Turbid, with thin, gray pellicle, 
and reddish-brown sediment. 

Litmus milk : Rapid reduction and 
proteolysis with odor of putrefaction. 

Potato : Echinulate, smooth, glistening, 
viscous, reddish-brown. 

Indole not produced. 

Nitrites are produced from nitrates. 

Acid from maltose and sucrose. No 
action on glucose, fructose, galactose, 
arabinose, lactose, raffinose, dextrin, 
inulin, salicin, amygdalin, glycerol. 

Ammonia is formed. 

Aerobic, facultative. 

Optimum temperature 21°C. 

Source : Isolated from tainted butter. 

Habitat : IVIilk, cream, butter, water, 
soil, creamery equipment (Long and 
Hammer, loc. cit. ; Claydon and Hammer, 



Iowa Agr. Exp. Sta., Res. Bull. 267, 
1939.) 

30. Pseudomonas mephitica Claydon 
and Hammer. (Jour. Bact., 37, 1939, 254.) 
From Latin, mephitis, pestilential, malo- 
dorous; M.L. Mephitis, a generic name. 

Rods : 0.5 to 1.0 by 1.5 to 14.0 microns, 
occurring singly, in pairs and in chains. 
Actively motile with a polar flagellum. 
Gram-negative. 

Gelatin: Slow liquefaction. 

Agar colonies: Convex, circular, about 
3 mm. in diameter, shiny, grayish white, 
edge entire, of the consistency of bread 
dough. 

Agar slant : Growth grayish-white, 
wrinkled, echinulate. After 1 or 2 days, 
a skunk-like odor develops. 

Broth: Turbid. Sediment. White 
pellicle. 

Potato: Growth echinulate, shiny, 
brownish. 

Litmus milk: A skunk-like odor devel- 
ops in 1 to 2 days. Grayish blue surface 
ring in about 3 days. Alkaline in 7 to 10 
daj's. In two weeks complete reduction. 
Slight proteoh^sis and viscosity. 

Hydrogen sulfide not produced. 

Indole not formed. 

Nitrites produced from nitrates. 

Acid but not gas produced slowly from 
glucose, fructose, maltose, and sucrose. 
No acid from arabinose, dextrin, galac- 
tose, glycerol, lactose, mannitol, raffinose 
or salicin. 
Aerobic, facultative. 
Optimum temperature 21°C. Growth 
slight at 5° and 30°C. No growth at 37°C. 
Source : Several cultures isolated from 
butter having a skunk-like odor. 
Habitat : Probably from water. 

31. Pseudomonas geniculata (Wright) 
Chester. {Bacillus geniculatus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 459; 
Chester, Man. Determ. Bact., 1901, 313; 
Achromobacter geniculatum Bergey et al.. 
Manual, 1st ed., 1923, 146.) From Latin, 



100 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



geniculatus, p.p. of geniculo, knotted, 
jointed. 

Medium-sized rods, occurring singly, 
in pairs and chains, motile, possessing 
polar flagella. Gram-negative. 

Gelatin colonies: Circular, whitish, 
translucent. Deep colonies yellowish. 

Gelatin stab : Infundibuliform liquefac- 
tion. Sediment light pink. 

Agar slant : Grayish, glistening, trans- 
lucent, limited, becoming brownish-graj'. 

Broth : Turbid, with slight gray pellicle 
and sediment. 

Litmus milk : Alkaline ; reduction of 
litmus; slight coagulation. 

Potato: Thin, brownish, moist, glis- 
tening, viscid. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 20° to 25°C. 

Habitat : Water. 

32. Pseudomonas fragi (Eichholz) 
Huss emend. Hussong, Long and Hammer. 
(Bacterium fragi Eichholz, Cent. f. 
Bakt., II Abt., 9, 1902, 425; Huss, Cent, 
f. Bakt., II Abt., 19, 1907, 661; Hussong, 
Long and Hammer, Iowa Agr. Exp. Sta. 
Res. Bull. 225, 1937, 122; also see Long 
and Hammer, Jour. Dairy Sci., 20, 1937, 
448.) From Latin /ra^um,. strawberry. 

Description from Hussong, Long and 
Hammer, loc. cit. 

Rods: 0.5 to 1.0 by 0.75 to 4.0 microns, 
occurring singly, in pairs and in chains. 
Motile with a polar flagellura. Gram- 
negative. 

Gelatin: Crateriform to stratiform 
liquefaction in 3 to 4 days. 

Agar colonies: Convex, glistening, gen- 
erally butyrous, occasionally viscid. 
Rough, smooth and intermediate forms 
are recognized in the description quoted. 
The rough forms are less proteolytic, and 
less active in the hydrolysis of fats. 

Agar slant : Growth abundant, spread- 
ing, raised, white, shiny, generally 
butyrous. Sweet ester-like odor resem- 
bling that of the flower of the May apple. 

Broth : Turbidity and sediment with a 
thin pellicle. 



Litmus milk : Acid ring followed by 
acid coagulum at surface. Complete 
coagulation in 2 to 3 weeks, some diges- 
tion. Characteristic May apple or straw- 
berry odor. 

Potato : Growth echiuulate to arbores- 
cent, raised, glistening, white, becoming 
brownish. 

Indole not produced. 

Nitrites not produced from nitrates. 

Ammonia produced from peptone. 

Hydrogen sulfide not produced. 

Acid from glucose and galactose, 
sometimes arabinose. No acid from 
glycerol, inulin, lactose, fructose, mal- 
tose, mannitol, raflfinose, salicin and 
sucrose. 

No acetylmethylcarbinol produced. 

Fat is generally hydrolyzed. 

Aerobic. 

Grows from 10° to 30°C. No growth at 
37°C. Very sensitive to heat. 

Source: Isolated from milk and other 
dairy products, dairy utensils, water, etc. 

Habitat : Soil and water. Widely dis- 
tributed (Morrison and Hammer, Jour. 
Dairy Sci., U, 1941, 9). 

Hussong (Thesis, Iowa State College, 
1932) regards Bacterium fragi Eichholz 
(loc. cit.) as the R type, Pseudomonas 
fragariae I Gruber (Cent. f. Bakt., II 
.\bt., 9, 1902, 705) as the O form, and 
Pseudomonas fragariae II Gruber (Cent, 
f. Bakt., II Abt., 14, 1905, 122) as the S 
form of the same organism. He makes no 
mention of Pseudomonas fragaroidea 
Huss (loc. cit.) which from its description 
would belong to the smooth type. A 
brief characterization of each of these 
organisms follows : (1 ) Bacterium fragi 
came from milk as drawn from an indi- 
vidual cow; it does not liquefy gelatin, 
exhibits no fluorescence, is strongly alka- 
line in litmus milk, and does not grow 
at 37°C, (2) Pseudomonas fragariae I 
came from fodder beets; it does not 
liquefy gelatin, has weak blue-greenish 
fluorescence, is weakly alkaline in milk, 
and grows at 37°C, (3) Pseudomonas 
fragariae II came from pasteurized milk ; 
it liquefies gelatin, coagulates milk, and 
does not grow at 37''C, (4) Pseudomonas 



FAMILY P8EUDOMONADACEAE 



101 



fragaroidea came from butter; it liquefies 
gelatin, coagulates milk, and grows at 

•33. Pseudomonas nebulosa (Wright) 
Chester. {Bacillus nebulosus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 465; 
Chester, Man. Determ. Bact., 1901, 311; 
Achromobacter nehulosum Bergey et al., 
Manual, 1st ed., 1923, 145; not Bacillus 
nebulosus Halle, These de Paris, 1898; 
not Bacillus nebulosus Vincent, Ann. 
Inst. Past., 21, 1907, 69; not Bacillus 
nebulosus Migula, Syst. d. Bakt., 2, 
1900, 844; not Bacillus nebulosus Gores- 
line, Jour. Bact., 27, 1934, 52.) From 
Latin, nebula, mist. 

Medium-sized rods, occurring singly. 
Motile, possessing polar flagella. Gram- 
negative. 

Gelatin colonies: Thin, circular, gray, 
translucent, hazy, with white center. 

Gelatin stab: Crateriform liquefaction. 

Agar slant: Thin, transparent streak. 

Broth: Turbid, with graj' sediment. 

Litmus milk: Alkaline; reduction of 
litmus. 

Potato: Scanty growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Sugar gelatin in deep stab : Fair growth, 
with some gas formation. 

Aerobic, facultative. 

Optimum temperature 30° to 35°C. 

Habitat : Water. 

Probable synonym : Pseudomonas cen- 
Irifugans Chester. (Man. Determ. 
Bact., 1901, 312; Bacillus centrifugans 
Wright, Mem. Nat. Acad. Sci., 7, 1895, 
462.) 

34. Pseudomonas coadunata (Wright) 
Chester. (Bacillus coadunatus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 460; 
Chester, iVIan. Determ. Bact., 1901, 310; 
Achromobacter coadunatum, Bergey et al., 
^Lanual, 1923, 147.) 1st ed.. From Latin, 
coadunatus, to unite closely. 



Medium-sized rods, with rounded 
ends, occurring singly, in pairs and in 
chains. Motile, possessing a polar flagel- 
lum. Gram -negative. 

Gelatin colonies: Circular, brownish, 
dense. 

Gelatin stab : Crateriform to stratiform 
liquefaction. 

Agar slant : Gray, translucent, 
spreading. 

Broth: Turbid, with gray pellicle and 
sediment. The medium has a slight 
greenish tint. 

Litmus milk: Acid; coagulated. 

Indole is formed. 

Nitrites not produced from nitrates. 

Sugar gelatin in deep stab : Good growth 
of discreet and confluent whitish colonies. 
Marked gas production; no liquefaction. 

Aerobic, facultative. 

Optimum temperature 20° to 25°C. 

Habitat : Water. 

35. Pseudomonas multistriata (Wright) 
Chester. {Bacillus multistriaius\N right. 
Memoirs Nat. Acad. Sci., 7, 1895, 462; 
Chester, Man. Determ. Bact., 1901, 310; 
Achromobacter multistrialum Bergey et 
al.. Manual, 1st ed., 1923, 147.) From 
Latin, multus, many, much; striatus, 
grooved. 

Medium-sized rods, with rounded ends, 
occurring singly and in pairs. Motile, 
possessing polar flagella. Gram-negative. 

Gelatin colonies : Circular, grayish- 
white, translucent. 

Gelatin stab : Crateriform liquefaction. 

Agar slant : Narrow, translucent, gray- 
ish streak. 

Broth: Turbid. 

Litmus milk : Slightly acid; coagulated. 

Potato : Grayish to creamy, thick, glis- 
tening, viscid, spreading. 

Indole not formed. 

Nitrites not produced from nitrates. 

Sugar gelatin in deep stab : Vigorous 
growth with marked gas production ; also 
liquefaction. 



* Prof. E. R. Hitchner, Univ. of Maine, Orono, Maine assisted in rearranging the 
descriptions of the acid and gas producing pseudomouads {Aeromonas) , April, 1943. 



102 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Aerobic, facultative. 

Optimum temperature 20° to 25°C. 

Habitat : Water. 

36. Pseudomonas punctata (Zimmer- 
mann) Chester. (Bacillus punctatus 
Zimmermann, Bakt. unserer Trink- unci 
Nutzwasser, Chemnitz, 1, 1890, 38; 
Bacillus aquatilis communis Kruse, in 
Fliigge, Die Mikroorganismen, 2, 1896, 
315; Bacterium -punctatum Lehmann and 
Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 
238; Chester, Man. Determ. Bact., 1901, 
313; Achromobacter punctatum Bergey et 
al., ]VIanual, 1st ed., 1923, 147.) From 
Latin, punctus, a puncture, point; M. L. 
punctate, dotted. 

Rods: 0.7 by 1.0 to 1.5 micron, occurring 
singly, in pairs and in chains. Motile 
with a single polar flagellum. Gram- 
negative. 

Gelatin colonies: Small, circular, gray, 
erose to filamentous, punctiform. 

Gelatin stab: Crateriform liquefaction. 
No pellicle. 

Agar slant: Gray, smooth, filamentous. 

Broth: Turbid with delicate pellicle. 

Litmus milk: Acid; coagulated; pep- 
tonized. 

Potato : Brownish-yellow to brownish- 
red color. 

Indole is formed. 

Nitrites not produced from nitrates. 

Hydrogen sulfide is formed. 

Acid and gas from glucose. 

Aerobic, facultative. 

Optimum temperature 25° to 30°C. 

Source : Common in the Chemnitz 
tap water. 

Habitat : Cause of a hemorrhagic septi- 
cemia in carp {Cyprinus) (Schaperclaus, 
Ztschr. f. Fischerei, 28, 1930; Cent. f. 
Bakt., II Abt., 105, 1942, 49). 

37. Pseudomonas hydrophila (Ches- 
ter) comb. nov. {Bacillus hydrophilus 
fuscus Sanarelli, Cent. f. Bakt., 9, 1891, 
222; Bacterium hydrophilus fuscus Ches- 
ter, Delaware College Agr. Expt. Sta., 
9th Ann. Rept., 1897, 92; Bacillus hydro- 



philus Chester, Manual Determ. Bact., 
1901, 235; Bacterium hydrophilum Weldin 
and Levine, Bact. Abs., 7, 1923, 14; 
Proteus hydrophilus Bergey et al.. Man- 
ual, 1st ed., 1923, 211; Aeromonas hydro- 
phila Stanier, Jour. Bact., 46, 1943, 213.) 
From Greek, hydor, water, philus, loving; 
M. L. water-loving. 

It was reported by Russell, Jour. Amer. 
Med. Assoc, 30, 1898, 1442 and later by 
Emerson and Norris, Jour. Exper. Med., 
7, 1905, 32 who made a complete study of 
its properties and its pathogenic action. 

Weldin (Iowa State College Jour. Sci., 
1, 1927, 151) considers Bacillus ranicida 
Ernst (Beitrage z. path. Anat. u. z. 
Allgemein. Pathol., 8, 1890, 204; Bac- 
terium ranicida Chester, Ann. Rept. 
Del. Col. Agr. Exp. Sta., 9, 1897, 141) a 
possible synonym of Proteus hydrophilus. 

Rods: 0.6 by 1.3 microns, occurring 
singly and in chains. Motile, with a 
single polar flagellum (Kulp and Borden, 
Jour, of Bact., U, 1942, 673). Gram- 
negative. 

Gelatin colonies: Small, circular, gray, 
translucent, stippled. 

Gelatin stab: Napiform liquefaction. 

Agar colonies : Whitish, raised, moist, 
stippled. 

Agar slant : Thin, whitish, glassy, 
spreading, becoming yellowish. 

Broth : Turbid, with heavy pellicle. 

Litmus milk: Acid; coagulated; pep- 
tonized. 

Potato: Yellowish-brown, moist 
slightly raised. 

Indole is formed. 

Nitrites produced from nitrates. 

Acid and gas from glucose, maltose, 
sucrose and mannitol. No action on lac- 
tose. 

Gas ratio H.rCO. = 1:4.71. Methyl 
red negative, acetylmethylcarbinol posi- 
tive, indol negative, citrate positive 
(Speck and Stark, Jour. Bact., U, 1942, 
697). 

Aerobic, facultative. 

Optimum temperature 37°C. 

Pathogenic for frogs, salamanders, fish, 



t'AMILY PSEtTDOMONADACEAE 



103 



mice, guinea pigs and rabbits, causing 
hemorrhagic septicemia. 

Distinctive characters: Much like 
Pseudo/nonas punctata (Guthrie and 
Hitchner, Jour. Bact., 43, 1943, 52). 

Source : Isolated from frogs dead of 
septicemia (red leg). 

Habitat : Water and infected fresh 
water animals. 

38. Pseudomonas ichthyosmia (Ham- 
mer) comb. nov. (Bacillus ichthyosmius 
Hammer, Iowa Agr. Sta. Res. Bui. 38, 
1917; Escherichia ichthyosmia Bergey et 
al., Manual, 1st ed., 1923, 201; Proteus 
ichthyosmius Bergey et al.. Manual, 4th 
ed., 1934, 364.) From Greek, ichthys, a 
fish; osme, an odor. 

Rods: 0.6 to 0.8 by 1.0 to M microns, 
occur singh'. Motile with a single polar 
flageilum (Breed). Gram-negative. 

Gelatin stab: Liquefaction. 

Agar colonies: Small, white, becoming 
darker with age. 

Agar slant : Dirty white, viscid growth. 

Broth : Turbid with gray sediment. 

Litmus milk: Acid. Litmus reduced. 
Cultures have fishy odor. 

Potato : Thin, glistening layer. 

Indole is formed. 

Nitrites produced from nitrates. 

Acid and gas from glucose, fructose, 
galactose, maltose, sucrose, glycerol, 
salicin and mannitol. Lactose, dulcitol, 
rafhnose and inulin not fermented. 

Aerobic, facultative. 

Optimum temperature 20^C. 

Source : Isolated from can of evaporated 
milk having a fishy odor. 

Habitat: Not known. 

39. Pseudomonas ambigua (Wright) 
Chester. (Bacillus ambiguus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 439; 
Chester, Man. Determ. Bact., 1901, 308; 
Achromobacler ambiguum Bergey et al.. 
Manual, 1st ed., 1923, 148.) From Latin, 
ambiguus, doubtful, uncertain. 

Small rods, with rounded ends, occur- 
ring singly, in pairs and in chains. Mo- 



tile, possessing a polar flageilum. Gram- 
negative. 

Gelatin colonies : Gray, translucent, 
slighth' raised, irregular, radiate, with 
transparent margin. 

Gelatin stab: No liquefaction. 

Agar slant : Gray, limited, entire. 

Broth: Turbid, with gray sediment. 

Litmus milk: Acid, slowly coagulated. 

Potato : Gray to creamy, viscid, 
spreading. 

Indole is formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 30° to 35°C. 

Habitat: Water. 

40. Pseudomonas sinuosa (Wright) 
Chester. (Bacillus sinuosus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 440; 
Chester, Man. Determ. Bact., 1901, 307; 
Achromobacler sinosum (sic) Bergey et 
al.. Manual, 1st ed., 1923, 148.) From 
Latin, sinuosus, full of bends, sinuous. 

Medium-sized rods, with rounded ends, 
occurring singly, in pairs and in chains, 
^lotile, possessing two to four polar 
flagella. Gram-negative. 

Gelatin colonies : Thin, translucent, 
irregular, center brownish. 

Gelatin stab: Grayish-white, glisten- 
ing, translucent. No liquefaction. 

Agar slant : Scanty, grayish growth. 

Broth : Turbid, with gray sediment. 

Litmus milk: Unchanged. 

Potato : Grayish-white, moist, spread- 
ing. 

Indole is formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 30° to 35°C. 

Habitat : Water. 

41. Pseudomonas cruciviae Gray^ and 
Thornton. (Gray and Thornton, Cent, 
f. Bakt., II Abt., 73, 1928, 91; Achro- 
mobacter cruciviae Bergey et al.. Manual, 
3rd ed., 1930, 218.) From Latin, crux, 
a cross, via, way, road; from Waycross, 
a place name. 



104 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Rods: 1.0 by 1.0 to 3.0 microns, occur- 
ring singly and in pairs. Motile with one 
to five polar flagella. Gram-negative. 

Gelatin colonies: Circular, white with 
buff center, convex, smooth, undulate. 

Gelatin stab : No liquefaction. 

Agar colonies : Circular or amoeboid, 
white to buff, flat to convex, smooth, 
entire. 

Agar slant: Filiform, pale buff, raised, 
smooth, undulate. 

Broth : Turbid. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

No acid in carbohydrate media. 

Attack phenol and m-cresol. 

Aerobic, facultative. 

Optimum temperature 30 to 35°C. 

Habitat : Soil. 

42. Pseudomonas rugosa (Wright) 
Chester. {Bacillus rugosus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 438; 
not Bacillus rugosus Henrici, Arb. Bakt. 
Inst. Tech. Hochsch. Karlsruhe, 1, 1894, 
28; not Bacillus rugosus Chester, Deter- 
minative Bacteriology, 1901, 220; Ches- 
ter, Determinative Bacteriology, 1901, 
323.) From Latin, rugosus, wrinkled. 

Small rods, with rounded ends, occur- 
ring singly, in pairs and in chains. Mo- 
tile, possessing one to four polar flagella. 
Gram-negative. 

Gelatin colonies : Grayish, translucent, 
slightly raised, irregular, sinuous, radi- 
ately erose to entire. 

Gelatin stab: Dense, grayish-green, 
limited, wrinkled, reticulate surface 
growth. No liquefaction. 

Agar slant: Grayish-white, limited, 
slightly wrinkled, translucent. 

Broth: Turbid, with grajdsh pelUcle 
and sediment. 

Litmus milk: Acid, coagulated. 

Potato: Moist, glistening, brown. 

Indole is formed. 

Nitrites not produced from nitrates. 

Aerobic. 

Optimum temperature 30°C. 

Habitat: Water. 



43. Pseudomonas desmolyticum Gray 
and Thornton. (Gray and Thornton, 
Cent. f. Bakt., II Abt., 7S, 1928, 90; 
Achromobacter desmolyticum Bergey et 
al.. Manual, 3rd ed., 1930, 217.) From 
Greek desinos, bond, band; lytikos, able 
to dissolve. 

Rods : 0.7 to 0.8 by 2.0 to 3.0 microns, 
occurring singly and in pairs. Motile, 
with one to five polar flagella. Gram- 
negative. 

Gelatin colonies: Circular, gray to 
buff, raised or umbonate. Smooth, glis- 
tening, entire. 

Gelatin stab: No liquefaction. 

Agar colonies: Circular or amoeboid, 
whitish, flat or convex, smooth, translu- 
cent to opaque, entire. 

Agar slant: Filiform, pale buff, raised, 
smooth, undulate. 

Broth : Turbid. 

Nitrites produced from nitrates. 

Starch not hydrolyzed. 

Acid from glucose. 

Attack phenol and naphthalene. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: Soil. 

44. Pseudomonas rathonis Gray and 
Thornton. (Gray and Thornton, Cent, 
f. Bakt., II Abt., 73, 1928, 90; Achromo- 
bacter rathonis Bergey et al., Manual, 
3rd ed., 1930, 216.) From M. L. of 
Ratho Park (Edinburgh). 

Small rods: 0.5 to 1.0 by 1.0 to 3.0 
microns. Motile, with polar flagella. 
Gram-negative. 

Gelatin colonies: Circular, white, 
raised, smooth, glistening, undulate. 

Gelatin stab: No Uquefaction. 

Agar colonies: Circular, buff, flat, 
smooth, glistening, entire. 

Agar slant : Filiform, pale buff, convex, 
smooth, glistening, undulate. 

Broth : Turbid, with peUicle. 

Nitrites produced from nitrates. 

Starch hydrolyzed. 

Acid from glucose and glycerol. 



FAMILY PSEUDOMONADACEAE 



105 



Attack phenol and cresol at times, also 
naphthalene. 

Aerobic, facultative. 
Optimum temperature 25°C. 
Habitat: Manure and soil. 

45. Pseudomonas dacunhae Gray and 
Thornton. (Gray and Thornton, Cent, 
f. Bakt., II Abt., 73, 1928, 90; Achro- 
mobacter dacv7ihae Bergey et al., Manual, 
3rd ed., 1930, 217.) From M. L. from 
the Island of d'Acunha. 

Rods: 0.5 to 0.8 by 1.5 to 3.0 microns. 
Motile with one to six polar flagella. 
Gram-negative. 

Gelatin colonies : Circular, whitish, 
raised, smooth, glistening, undulate. 

Gelatin stab: Xo liquefaction. 

Agar colonies: Circular to amoeboid, 
white, flat, glistening, opaque, entire. 

Agar slant: Filiform, pale buff, raised, 
smooth, glistening, undulate. 

Broth: Turbid. 

Nitrites produced from nitrates. 

Starch not hydrolyzed. 

No acid from carbohydrate media. 

Attack phenol. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat : Soil . 

46. Pseudomonas arvilla Gray and 
Thornton. (Gray and Thornton, Cent, 
f. Bakt., II Abt., 73, 1928, 90; Achromo- 
bacter arvillum Bergey et al., Manual, 
3rd ed., 1930, 217.) From Latin, arvum, 
an arable field; M. L. dim. a little field. 

Rods: 0.5 to 0.7 by 2.0 to 3.0 microns. 
Motile with one to five polar flagella. 
Gram-negative. 

Gelatin colonies: Circular, whitish, 
convex, smooth, glistening, lobate. 

Gelatin stab: No liquefaction. 

Agar colonies: Circular or amoeboid, 
white to buff, flat to convex, smooth, 
glistening, opaque, entire. 

Agar slant : Filiform, whitish, concave, 
smooth, ringed, entire. 

Broth : Turbid. 

Nitrites not produced from nitrates. 



Starch not hydrolyzed. 
Acid from glucose. 
Attacks naphthalene. 
Aerobic, facultative. 
Optimum temperature 25°C. 
Habitat: Soil. 

47. Pseudomonas salopium Gray and 
Thornton. (Gray and Thornton, Cent, 
f. Bakt., II Abt., 73, 1928, 91; Achromo- 
bacter salopium Bergey et al.. Manual, 
3rd ed., 1930, 219.) From Latin, Salop, 
Shropshire. 

Rods: 0.7 to 1.0 by 1.0 to 3.0 microns, 
occurring singly and in pairs. Motile 
with one to six polar flagella. Gram- 
negative. 

Gelatin colonies: Circular, grayish- 
buff, flat, rugose or ringed, translucent 
border. 

Gelatin stab: No liquefaction. 

Agar colonies : Circular or amoeboid, 
white to buff, flat to convex, smooth, 
glistening, translucent border, entire. 

Agar slant: Filiform, whitish, raised, 
smooth, glistening, lobate. 

Broth: Turbid with pellicle. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Acid from glucose and sucrose. 

Attacks naphthalene. 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: Soil. 

48. Pseudomonas minuscula McBeth. 
(McBeth, Soil Science, 1, 1916, 437; 
Cellidomonas minuscula Bergey et al.. 
Manual, 1st ed., 1923, 162.) From Latin 
dim. rather small. 

Rods: 0.5 by 0.9 micron. Motile with 
one to two polar flagella. Gram-negative. 

Gelatin stab : Moderate growth . Slight 
napiform liquefaction. 

Agar colonies: Small, circular, slightly 
convex, butyrous becoming brittle, gray- 
ish-white, finely granular, entire. 

Agar slant: Moderate, flat, grayish- 
white. 

Broth : Turbid. 



106 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Litmus milk : Acid, not digested. 
Potato: No apparent growth. 
Indole is formed. 
Nitrites produced from nitrates. 
Ammonia is produced. 
Acid from glucose, lactose, maltose, 
sucrose and starch. 
Aerobic, facultative. 
Optimum temperature 20°C. 
Habitat : Soil. 

49. Pseudomonas tralucida Kellerman 
et al. (Kellerman, McBeth Scales and 
Smith, Cent. f. Bakt., II Abt., 39, 1913, 
37; Cellulomonas tralucida Bergey et al., 
Manual 1st ed., 1923, 163.) From Latin, 
clear, transparent. 

Rods : 0.6 by 1.2 microns. Motile with 
one or two polar flagella. Gram-negative. 

Gelatin stab : No liquefaction. 

Agar slant: Scant, grayish growth. 

Broth : Turbid. 

Litmus milk : Acid. 

Potato: No growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Ammonia not produced. 

Acid from glucose, maltose, lactose, 
sucrose, starch, glycerol and mannitol. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Habitat: Soil. 

50. Pseudomonas mira McBeth. (Mc- 
Beth, Soil Science, 1, 1916, 437; Cellu- 
lonwnas mira Bergey et al.. Manual, 1st 
ed., 1923, 165.) From Latin, minis, won- 
derful, extraordinary. 

Rods : 0.4 by 1.6 microns. Motile with 
a single polar flagellum. Gram-negative. 

Gelatin stab: Good growth. No lique- 
faction. 

Agar colonies : Circular, convex, gray- 
ish-white, granular, lacerate. 

Agar slant: Moderate, flat, grayish- 
white, somewhat iridescent. 

Broth: Turbid. 

Litmus milk: Alkaline. 

Potato: Moderate, grayish-white. 

Indole not formed. 



Nitrites produced from nitrates. 
Ammonia is produced. 
Acid from glucose, maltose, lactose, 
sucrose, starch, glycerol and mannitol. 
Aerobic, facultative. 
Optimum temperature 20°C. 
Habitat: Soil. 

51. Pseudomonas lindneri Kluyver 
and Hoppenbrouwers. (Lindner, 50 Ju- 
bilaumsber. Westpreuss. Bot.-Zool. 
Vereins, 1928, 253; T ermohacterium mo- 
bile Lindner, Atlas d. Mikrosk. Grundl. 
d. Garungsk., 3 Aufl., 2, 1928, Taf. 68; 
Kluyver and Hoppenbrouwers, Arch. f. 
Mikrobiol., 2, 1931, 259; Achromobacter 
mobile Kluyver and Hoppenbrouwers, 
ibid., 258; not Pseudomonas mobilis 
Migula, Syst. d. Bakt., 2, 1900, 923.) 
Named for Lindner, the German bacteri- 
ologist who first studied this organism. 

Short rods 1.4 to 2.0 by 4.0 to 5.0 mi- 
crons. Occurring singly, in pairs and 
short chains. Motile with a single polar 
flagellum. Gram-negative. 

Peptone gelatin: Poor growth. 

Peptone agar: Poor growth. 

Wort agar: White, round, raised colo- 
nies, 1 mm. in diameter. Good growth. 
Still better where 2 per cent sucrose, or 
yeast extract with sucrose is added. 
Chalk added to neutralize acid. 

Broth : Poor growth in peptone or yeast 
extract broth unless sugars are added. 

Carbon dioxide, ethyl alcohol and some 
lactic acid produced from glucose and 
fructose, but not from mannose. May 
or may not ferment sucrose. May pro- 
duce as much as 10 per cent alcohol. 

Catalase produced. 

Anaerobic, facultative. 

Optimum temperature 30°C. 

Distinctive character: The fermenta- 
tion resembles the alcoholic fermentation 
produced by yeasts. 

Source : Isolated from the fermenting 
sap (pulque) of Agave americana in 
Mexico. 

Habitat: Fermenting plant juices in 
tropical countries (Mexico). 



FAMILY PSEUDOMOXADACEAE 



107 



52. Pseudomonas membranoformis 
(Zobell and Allen) Zobell. (Achromo- 
bacter membranoformis Zobell and Allen, 
Jour. Bact., 29, 1935, 246; Zobell, Jour. 
Bact., 46, 1943, 45). From Latin jnem- 
brana, membrane, and forma having the 
form of. 

Rods: 0.9 to 1.2 by 3.5 to 4.8 microns, 
occurring singly and in pairs. Motile 
with lophotrichous flagella. Encapsu- 
lated. Gram-negative. 

Gelatin stab: Growth fihform, best at 
top, with slow crateriform licjuefaction. 

Agar colonies: Circular, 1.0 to 2.5 mm, 
with crinkled surface. 

Agar slant : Moderate, beaded, raised 
growth. Membranous consistency. Be- 
comes browned with age. 

Broth : Slight turbidity, tl(jcculent sedi- 
ment, film of growth on walls of test tube. 

iNIilk: Xo growth. 

Potato: Xo growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

No H2S produced. 

Acid but not gas from glucose, sucrose, 
dextrin and mannitol. Xo acid from 
lactose or xylose. 

Xo diastatic action. 

Optimum temperature 20° to 25°C. 

Aerobic. 

Source : Sea water. 

Habitat : Sea water. 

53. Pseudomonas marinoglutinosa 

(Zobell and Allen) Zobell. (Achromo- 
bacter marinoglutinosus Zobell and Allen, 
Jour. Bact., 29, 1935, 246; Zobell, Jour. 
Bact., 46, 1943, 45). From Latin 
marinas, pertaining to the sea, and 
glutinosus, full of glue, sticky. 

Short rods: 0.7 to 1.0 by 1.8 to 2.4 mi- 
crons, with rounded ends, occurring 
singly, in pairs and in clumps. Motile 
with polar flagella. Staining granular. 
Encapsulated. Gram-negative. 

Gelatin stab : Moderate filiform growth 
with slight napiform liquefaction. Xo 
pigment. 

Agar colonies : Ptound with concentric 



circles and crinkled radial lines, 1.5 to 
5.0 mm. in diameter. No pigment. 

Agar slant: Moderate, filiform, flat. 
Butyrous consistency. 

Broth: Moderate clouding, marked 
ring, adherent film of growth on test tube 
wall, and flaky sediment. 

Milk : No growth. 

Potato: Xo growth. 

Indole not formed. 

Xitrites not produced from nitrates. 

Hydrogen sulfide and ammonia pro- 
duced from Bacto-tryptone. 

Acid but not gas from xylose and dex- 
trin. Xo acid from glucose, lactose, 
sucrose and mannitol. 

Starch is hydrolyzed. 

Optimum temperature 20° to 25°C. 

Aerobic, facultative. 

Source : Sea water. 

Habitat: Sea water. 

54. Pseudomonas gelatica (Gran) 
Bergey et al. (Bacillus gelaticus Gran, 
Bergens Museums Aarbog., 1902, 14; 
Bacterium gelaticum Lundestad, Cent, 
f. Bakt., II Abt., 75, 1928, 328; Bergey 
et al., Manual, 3rd ed., 1930, 175.) From 
French, like gelatin. 

Pods, ^vith rounded ends, 0.6 to 1.2 by 
1.2 to 2.6 microns, occurring singly, in 
pairs, and sometimes in short chains. 
Motile. Gram-negative. 

Fish-gelatin colonies : Circular, trans- 
parent, glistening, becoming brownish in 
color. 

Fish-gelatin stab : Liquefaction infun- 
dibuliform, with greenish color. 

Sea-weed agar colonies : Circular, flat, 
entire, glistening, reddish-brown center 
with gra.vish -white periphery. Lique- 
fied. 

Fish-agar slant: Flat, transparent 
streak, with undulate margin, reddish- 
brown . 

Broth : Turbid with flocculent pellicle, 
and greenish-yellow sediment. 

Indole not formed. 

Xitrites are produced from nitrates. 

Starch hydrolyzed. 



108 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



No action on sugars. 
Anaerobic, facultative. 
Optimum temperature 20 to 25°C. 
Habitat : Sea water of Norwegian coast. 

55. Pseudomonas calcis (Drew) 
Kellerman and Smith. {Bacterium cal- 
cis Drew, Yearbook Carnegie Inst. 
Wash., 11, 1912, 136-144; Kellerman and 
Smith, Proc. Nat. Acad. Sci., 4, 1914, 
400.) From Latin calx (calc-), lime. 

Ovoid rods, 1.1 by 1.5 to 3 microns, 
usually single but may form long chains. 
Actively motile with one polar flagellum. 
Gram-negative. 

Grows best in sea water or 3 per cent 
salt media. Deposits CaCOs. 

Agar colonies : Circular, with finely 
irregular outline, granular appearance, 
elevated, spreading; old colonies having 
brownish tinge in center. 

Gelatin stab : Infundibuliform lique- 
faction. 

Gelatin colonies: Small, with lique- 
faction. 

Broth : Good growth especially in pres- 
ence of potassium nitrate, peptone or 
calcium malate. 

Acid from glucose, mannite and sucrose 
but not from lactose. 

Nitrates reduced to nitrites and am- 
monia. 

Aerobic, facultative. 

Optimum temperature 20 to 28°C. 

Habitat : Sea water and marine mud. 

Bavendarmn (Arch. f. Mikrobiol., 3, 
1932, 214) states that Pseudomonas calcis 
is probably synonymous with Bacterium 
brandti, Bacterium bauri and Bacterium 
feiteli described by Parlandt (Bull. 
Jard. imp. Bot. St. Petersburg, 11, 1911, 
97-105). 

56. Pseudomonas calciprecipitans Mo- 
lisch. (Cent. f. Bakt., II Abt., 65, 1925, 
130.) From Latin, calx (calc-), lime; 
praecipito, to cast down headlong, to 
precipitate. 

Thin rods: 0.5 to 0.8 by 1.5 to 3.6 mi- 
crons, with rounded ends, often staining 



irregularly. Motile, with one polar flagel- 
lum. Gram -negative. 

Gelatin colonies : Circular, light brown 
in color (large colonies show CaCOs 
crystals). 

Gelatin stab : Surface growth with fili- 
form growth in depth. Liquefaction 
starts at bottom. 

Agar colonies (sea water). Grayish- 
white, glistening. In two to three weeks 
crystals of calcium carbonate form in the 
agar. 

Agar slant: Slight, whitish, surface 
growth, becoming thick, spreading, glis- 
tening, with abundant CaCOs crystals in 
medium. 

Ammonia formed. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Habitat : Sea water. 

57. Pseudomonas ichthyodermis (Wells 
Zobell) Zobell and Upham. (Achromo- 
bacter ichthyodermis Wells and Zobell, 
Proc. Nat. Acad. Sci., 20, 123, 1934; 
Zobell and Upham, Bull. Scripps Inst. 
Oceanography, S, 1944, 246 and 253.) 
From Greek, ichthys, a fish; derma, sldn. 

Small rods, 0.9 to 1.3 by 3 to 5 microns, 
occurring singly and in pairs. No spores. 
Encapsulated. Polar flagella. Pleomor- 
phic forms predominate in old cultures. 
Gram-negative. 

Requires sea water following initial 
isolation. The following differential me- 
dia are prepared with sea water. 

Agar colonies : Glistening, colorless, 
convex, circular colonies 2 to 4 mm. in 
diameter. 

Agar slants : Abundant, filiform, raised, 
smooth, opalescent growth. 

Gelatin tube : Rapid crateriform lique- 
faction complete in 5 days at 18°C. 

Sea water broth : Turbidity, with pelli- 
cle, little granular sediment and no odor. 

Milk: No growth. Casein digested 
when 3 per cent salt is added. 

Potato : No growth unless dialyzed in 
sea water. Then fair growth with nc 
pigment. 



FAMILY PSEUDOMONADACEAE 



109 



Acid from glucose, maltose, sucrose 
and mannitol but not from lactose or 
glycerol. 

Starch hydrolyzed. 

Ammonia liberated from peptone but 
.no hydrogen sulfide produced. 

Indole formed in tryptophane sea water 
broth. 

Nitrites produced from nitrates. 

Optimum temperature 20 to 25°C.; 
30°C. incubation will kill recently isolated 
organisms. 

Aerobic, facultative. 

Source : Isolated from diseased kilifish 
(Fundulvs parvipinnis) . 

Habitat : Skin lesions and muscle tissue 
of infected marine fish. 

58. Pseudomonas nigrifaciens White. 
(Scientific Agriculture, W, 1940, 643.) 
From Latin viger, black and faciens, 
making. 

Rods: 0.3 to 0.7 by 1 to 5 microns, oc- 
curring singly or in pairs, and having 
rounded ends. Actively motile, with a 
single polar flagellum. Gram-negative. 

Gelatin stab : Pigmented surface growth 
after 24 hours. Slight crateriform lique- 
faction changing to saccate. 

Agar colonies: Circular, convex, 
smooth, glistening, entire, 2 to 4 mm in 
diameter. Slight fluorescence in earl}- 
stages. The medium assumes a brownish 
color. 

Agar slant : Growth filiform, smooth, 
moist, glistening, with blackish pigmenta- 
tion at 4° and 15°C. in 48 hrs., the medium 
turning brownish. Slight fluorescence in 
early stages. 

Broth: Turbid after 24 hours. .A.fter 
5 to 6 days a black ring and then a pellicle 
forms, later a black sediment. Medium 
turns brown. 

Litmus milk : A black ring appears 
after 3 days at 15°C. followed by a pellicle. 
Litmus is reduced. Alkaline reaction. 
No coagulation. Digested with a putrid 
odor. 

Indole not formed. 



Nitrites not produced from nitrates in 
7 days. No gas produced. 

iStarch is hydroh'zed. Natural fats 
not hydrolyzed. 

Alkaline reaction produced in sucrose, 
maltose, lactose, glucose, mannitol and 
raffinose broth (pH 8.2). No gas pro- 
duced. 

Ammonia produced in peptone broth. 

Aerobic. 

Optimum pH 6.8 to 8.4. 

Temperature relations: Minimum 4°C. 
Optimum 25°C. Ma.ximum 33-35°C. 

Distinctive characters : No or slow 
growth in culture media in the absence of 
salt. Maximum growth and pigmenta- 
tion appeared with 1.5 and 2.5 per cent 
salt. Optimum pigmentation occurs at 
4" and 15°C. 

Source : Several cultures isolated from 
samples of discolored butter. 

Habitat : Causes a black to reddish- 
brown discoloration of print butter. 
Evidently widely distributed in nature. 

59. Pseudomonas beijerinckii Hof. 

(Travaux botaniques neerlandais, S2, 
1935, 152.) Named for M. W. Beijerinck, 
Dutch bacteriologist . 

Small rods: Motile with polar flagella. 

Gelatin: No liquefaction. 

Indole not formed. 

Nitrites produced from nitrates by 
four out of six strains. 

Cellulose not decomposed. 

Acid from glucose. In yeast-water 
with 2 per cent glucose and 12 per cent 
NaCl no gas is produced. 

Pigment production : Insoluble purple 
pigment produced but not in all media; 
is localized markedly; reduced oxygen 
tension necessary; optimum pH 8.0; not 
produced in yeast-water or in peptone- 
water; produced only when grown in 
extracts of beans or some other vegetable. 

Aerobic. 

Source : Six strains isolated from beans 
preserved with salt. 

Habitat: Causes purple discoloration 
of salted beans. 



110 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



60. Pseudomonas salinaria Harrison 
and Kennedy. (Harrison and Kennedy, 
Trans. Royal Soc. of Canada, 16, 1922, 
121; Serratia salinaria Bergey et al.. 
Manual, 1st ed., 1923, 93; Flavobacterium 
(Halobacterium) salinarium Elazari-Vol- 
cani. Studies on the Microflora of the 
Dead Sea, Thesis, Hebrew Univ., Jeru- 
salem, 1940, 59.) From Latin, salinae, 
saltworks. 

Probable synonym : Serratia sambhari- 
anus Dixit and Vachna, Current Sci., 11, 
1942, 107 (see Biol. Abs., 17, 1943, 793). 
Halophilic growing in 30 per cent salt. 
From salt lake in India. 

Occurs as spheres and rods, 2.0 to 3.0 
microns in diameter, 1.0 to 1.6 by 3.0 
to 15.0 microns, occurring singly, as 
ovoid, amoeboid, clavate, cuneate, trun- 
cate, spindle, club, pear-shape, and 
irregular forms. Motile, frequently with 
a fiagellum at each pole. Gram-negative. 

Does not grow on ordinary culture 
media. Grows well on salted fish. 

Codfish agar (16 to 30 per cent salt) : 
Growth slow, smooth, raised, coarsely 
granular, entire, pale pink to scarlet 
(Ridgway chart). 

No acid from carbohydrate media. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 42°C. 

Source : Isolated from cured codfish 
(Harrison and Kennedy, loc. cit.). Iso- 
lated from salted fish by Browne (Absts. 
Bact., 6, 1922, 25 and Proc. Soc. Exper. 
Biol, and Med., 19, 1922, 321) who re- 
garded this pleomorphic bacterium as 
two organisms — Spirochaeta halophiiica 
and Bacterium halophiiica. 

Habitat: Produces reddening of dried 
codfish and causes rusty herring. In 
sea salt, and salt ponds containing not 
less than 16 per cent salt. 

61. Pseudomonas cutirubra Lochhead. 
{Serratia cutirubra Lochhead, Can. Jour. 



of Research, 10, 1934, 275; Bacterium 
cutirubrum Lochhead, Jour. Bact., 27, 
1934, 62; ibid., J^5, 1943, 575; Flavobac- 
terium {Halobacterium cutirubrum Elaz- 
ari-Volcani, loc. cit., 59.) From Latin, 
cutis, skin, hide; ruber, red. 

Occurs as rods and spheres. Spheres 
1 to 1.5 microns in diameter. Rods 1.5 
to 8.0 by 0.7-1.4 microns. Rod forms 
motile with single polar flagellum. Coc- 
coid forms motile when young. Gram- 
negative. 

No growth on ordinary media. 

Milk agar (20 per cent salt to satura- 
tion; optimum 28-32 per cent) : Colonies 
3-4 mm. in diameter, round and slightly 
convex. Pink to rose doree (Ridgway 
chart). 

Milk agar slants: Growth filiform, 
slightly spreading, rather flat with 
smooth, glistening surface and mem- 
branous consistency. Proteolytic action. 

Liquid media: No or slight growth. 

Gelatin (salt ) : Pronounced liquefaction . 

Indole not formed, Lochhead {loc. cit.). 
Faint test. Gibbons (Jour. Biol. Board 
Canada, 3, 1936, 75). 

Nitrites not produced from nitrates. 

Diastatic action negative. 

No carbohydrate fermentation. 

Aerobic, obligate. 

Optimum temperature 37°C. 

Halophilic. 

Source : Isolated from salted hides. 

Habitat: Sea water and sea salt. 

62. Pseudomonas harveyi Johnson 
comb. nov. {Achromobacter harveyi John- 
son and Shunk, Jour. Bact., 31, 1936, 
587.) Named for E. N. Harvey, who 
studied luminous bacteria. 

Rods: 0.5 to 1.0 by 1.2 to 2.5 microns, 
occurring singly or in pairs, with rounded 
ends. Occasionally slightly curved; ends 
occasionally slightly pointed. Non-spore- 
forming. Capsules absent. Motile with 
a single polar flagellum, 2 to 3 times the 
length of the cell. Gram-negative. 



FAMILY PSEUDOMONADACEAE 



111 



Sea water gelatin colonies: After 24 
hours at 20°C, circular, about 1.5 mm. in 
diameter or larger, margin slightly undu- 
late, sunken due to the beginning of 
liquefaction, interior somewhat zonate; 
colonies surrounded bj' a halo of numerous 
small secondarj' colonies, circular and 
finely granular. In crowded plates a 
large number of gas bubbles are formed. 
Luminescent. 

Sea water gelatin stab : Rapid saccate 
liquefaction complete in 5 days at 22°C. 
Abundant flocculent sediment. 

Sea water agar colonies : Mostly very 
large, 6 to 8 cm. in diameter in 24 hours, 
flat, highly iridescent, circular with 
undulate margin, or composed of narrow 
and close or wide filamentous growth. 
Occasionally small colonies appear that 
are circular, with entire or slightly undu- 
late margin, often producing irregular 
secondary growth, surface always smooth. 
Luminescent. 

Sea water agar slant : Growth abundant, 
spreading, grayishly viscous, homogene- 
ous, iridescent, the medium becoming 
rapidly alkaline when inoculated at an 
initial pH of 7.0. With fish decoctions 
added to the medium, luminescence is 
much brighter and growth becomes 
brownish after several days. 

Growth on autoclaved fish: Abundant, 
smooth, glistening, yellowish, becoming 
dirty brown after several days. Mild 
putrefactive odor. Luminescence very 
brilliant. 

Sea water containing 0.2 per cent 
peptone: Abundant uniform turbidity, 
thin pellicle, sediment accumulating over 
a period of several days. Luminescence 
at surface only unless the tube is shaken. 

Milk, with or without the addition of 
2.8 per cent salt: Xo growth. 

Potato plugs resting on cotton saturated 
with sea water: Growth slight, somewhat 
spreading, slightlj^ brownish. Luminous. 

Indole is formed (Gore's method). 

Nitrites are produced from nitrates. 

Ammonia is produced in peptone media 
(Hansen method). 



Fixed acid from glucose, fructose, 
mannose, galactose, sucrose, maltose, 
mannitol, dextrin, glycogen, trehalose, 
cellobiose; slowly from salicin. Xon- 
fixed acid from melezitose; slight acid 
from sorbitol, disappearing in 24 hours. 
No acid from gh'cerol, xylose, arabinose, 
dulcitol, inositol, adonitol, erythritol, 
arabitol, lactose, raffinose, rhamnose, 
fucose or alpha methyl glucoside. 

Starch agar: Wide zone of hydrolysis. 

Hj'drogen sulfide is produced (Zobell 
and Fantham method). 

Temperature relations : Optimum 35° 
to 39°C. Abundant growth at 22° to 
25°C. Optimum luminescence at 20° to 
40°C. 

Not pathogenic for white rats or amphi- 
pods. 

Aerobic, facultative anaerobe. 

Source : Isolated from a dead amphipod 
(Talorchestia sp.) at Woods Hole, Massa- 
chusetts. 

Habitat : Sea water. 

63. Pseudomonas phosphorescens 

(Fischer) Bergey et al. (Bacillus phos- 
phorescens Fischer, Zeitschr. f. Hyg., 2, 
1887, 58; Photobacterium indicum Bei- 
jerinck, Arch. Neerl. d. Sci. Exactes, 23, 
1889, 401 ; Bacterium phosphorescens 
Lehmann, Cent. f. Bakt., 5, 1889, 785; 
not Bacterium phosphorescens Fischer, 
Cent. f. Bakt., 3, 1888, 107; Bergey et al., 
Manual, 3rd ed., 1930, 177.) From Greek 
phosphoreo, to bear or bring light. 

See page 699 for additional synonjins. 

Description taken from Fischer {loc. 
cit.). 

Small, thick rods: 2 to 3 times as long 
as wide, with rounded ends. Motile. 
Stain lightly with aniline dj'es. 

Gelatin colonies: After 36 hours, small, 
circular, gray-white, punctiform. 
Liquefaction. Bluish to green phos- 
phorescence in 4 to 5 days. 

Blood serum: Gray-white, slimy 
growth. 

Potato: Thin white layer in 2 to 3 
days. 



112 



MANUAL OF DETERMINATIVE BACTERIOLOGt 



Cooked tish: Abundant growth. En- 
tire surface covered with a gray -white, 
slimy growth. Bluish-white phos- 
phorescence. 

Alkaline broth: Slight turbidity in 24 
hours. Pellicle in 3 days. 

Acid broth: No turbidity. No phos- 
phorescence. 

Milk: No growth. 

No gas formed. 

Not pathogenic for Iaborator.y 
animals. 

Aerobic. 

Optimum temperature 20° to 30°C. 

Source: From sea water of the West 
Indies. 

Habitat: Sea water. 

64. Pseudomonas pierantonii (Zirpolo) 
Bergey et al. (Micrococcus pierantonii 
Zirpolo, Boll. del. Societa dei Natural, 
in Napoli, SI, 1918, 75; Cocco -bacillus 
pierantonii Meissner, Cent. f. Bakt., II 
Abt., 67, 1926, 204; Bergey et al., Manual, 
3rd ed., 1930, 176.) Named for Fieran- 
toni, an Italian. 

Oval rods: 0.8 by 1.0 to 2.0 microns. 
Polymorphic rods, sometimes vacuolated. 
Motile. Gram-negative. 

Gelatin colonies: Circular, luminous. 

Gelatin stab: Not liquefied. 

Sepia agar colonies: Circular, white, 
convex, smooth, serrate edge. Intense 
greenish luminescence. 

Egg-glycerol agar slant : Yellowish- 
green, luminous streak. 

Broth: Turbid. 

Indole not formed. 

Acid from glucose and maltose, some 
strains also produce acid from lactose and 
sucrose. 

Best growth in alkaline media. 

Aerobic. 

Optimum temperature 33°C. 

Source: Isolated from the photogenic 
organ of the cephalopod Rondeletia minor. 



*65. Pseudomonas martyniae (Elliott) 
Stapp. (Bacterium martyniae Elliott, 
Jour. Agr. Res., 29, 1924, 490; Stapp, in 
Sorauer, Handbuch der Pflanzenkr., 2, 
5 Auf., 1928, 278; Phytomonas martyniae 
Bergey et al.. Manual, 3rd ed., 1930, 262.) 
From M. L. Martynia, a generic name. 

Rods: 0.59 to 1.68 microns. Capsules. 
Chains. Motile with one to several 
bipolar flagella. Gram-negative. 

Green fluorescent pigment produced. 

Gelatin: Liquefied. 

Beef agar colonies: White, round, 
smooth, glistening, raised. 

Broth : Clouding in bands. Thin pelli- 
cle. Small crystals. 

Milk : Soft acid curd with peptoniza- 
tion. 

Nitrites produced from nitrates after 2 
weeks. 

Indole not produced. 

Hydrogen sulfide production slight. 

Acid but not gas from glucose, galac- 
tose, arabinose and sucrose. No acid 
from rhamnose, lactose, maltose, raffinose, 
mannitol and glycerol. 

Starch hydrolysis none or feeble. 

Optimum temperature 26°C. Maxi- 
mum 37°C. Minimum 1.5°C. 

Optimum pH 6.0 to 6.7. pH range 5.4 
to 8.9 . 

Aerobic. 

Source : Isolated from diseased leaves 
of the unicorn plant from Kansas. 

Habitat : Pathogenic on Martynia 
louisiana. 

66. Pseudomonas striafaciens (Elli- 
ott) Burkholder. (Bacterium stria- 
faciens Elliott, Jour. Agr. Res., 35, 1927, 
823; Phytomonas striafaciens Bergey 
et al., Manual, 3rd ed., 1930, 268; Burk- 
holder, Phytopath., 32, 1942, 601.) 
From L. stria, stripe; faciens, making. 



* The section covering the pseudomonads that cause plant diseases has been re- 
vised by Prof. Walter H. Burkholder, Cornell Univ., Ithaca, New York, April, 1943. 



FAMILY PSEUDOMONADACEAE 



113 



referring to the type of lesion caused 
on the blades of oats. 

Rods: 0.66 by 1.76 microns. Motile 
with one to several flagella. Capsules. 
Gram-negative. 

Green fluorescent pigment produced. 

Gelatin: Liquefied. 

Beef -peptone agar colonies : White, 
raised, margins entire or slightly undu- 
lating. 

Broth : Clouding in layers. Ring and 
slight pellicle. 

Milk: Alkaline, sometimes a soft curd 
which digests or clears. 

Slight production of nitrites from 
nitrates. 

Indole not produced. 

Acid but not gas from glucose, fructose 
and sucrose. No acid from lactose, mal- 
tose, glycerol and mannitol. 

Starch: Hydrolysis slight. 

Optimum temperature 22°C. 

Optimum pH 6.5 to 7.0. 

Aerobic. 

Distinctive characters: Differs from 
Pseudomonas coronafaciens in that the 
cells are somewhat smaller and the patho- 
gen produces a streak on oat blades 
instead of a halo spot. 

Source : Forty cultures isolated from 
oats gathered in various parts of America. 

Habitat : Pathogenic on cultivated oats, 
and to a slight degree, on barley. 

67. Pseudomonas tomato (Okabe) 
comb. 710V. {Bacterium tomato Okabe, 
Jour. Soc. Trop. Agr. Formosa, 5, 1933, 32; 
Phytomonas tomato Magrou, in Hauduroy 
et al.. Diet. d. Bact. Path., Paris, 1937, 
422.) Named for the host plant, tomato. 

Probable synonym : Bacterium punc- 
tidans Bryan, Phytopath., 23, 1933, 897. 

Rods : 0.69 to 0.97 by 1.8 to 6.8 microns. 
Motile with 1 to 3 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Slow liquefaction. 

Beef -extract agar colonies: White, 
circular, flat and glistening. 

Broth: Turbid in 24 hours. Pellicle. 



Milk: Becomes alkaline and clears. 

Nitrites are usually produced from 
nitrates. 

Indole not produced. 

No H2S produced. 

Acid but not gas from glucose, sucrose 
and lactose. No acid from maltose and 
glycerol. 

Starch hydrolysis feeble. 

Slight growth in 3 per cent salt. 

Optimum temperature 20° to 25°C. 
Ma.ximum 33°C. 

Aerobic. 

Source : Isolated from diseased tomato 
leaves. 

Habitat : Pathogenic on tomato, Lyco- 
persicon esculentum. 

68. Pseudomonas aceris (Ark) Burk- 
holder. {Phjitomonas aceris Ark, Phyto- 
path., £9, 1939, 969; Burkholder, Phyto- 
path., 32, 1942, 601.) From Latin acer, 
maple; M.L. Acer, generic name. 

Rods: 0.3 to 0.8 by 0.8 to 2.5 microns. 
Motile with 1 to 2 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced. 

Gelatin: Liquified. 

Beef -extract-peptone agar : Colonies are 
grayish-white. Appearing in 24 hours. 

Broth: Turbid. 

Milk: Clearing with no coagulation. 

Nitrites not produced from nitrates. 

Indole not produced. 

Hydrogen sulfide not produced. 

Acid from glucose, fructose, galac- 
tose, arabinose, xylose, sucrose, maltose, 
lactose, raffinose, mannitol, glycerol and 
dulcitol. 

Slight growth in broth plus 6 per cent 
salt (Burkholder). 

Temperature : 13° to 3rC. 

Source : From diseased leaves of the 
large leaf maple, Acer macrophillum . 

Habitat: Causes a disease of Acer spp. 

69. Pseudomonas angulata (Fromme 
and Murray) Holland. (Bacterium an- 
gulatum Fromme and Murray, Jour. Agr. 
Res., 16, 1919, 219; Holland, Jour. Bact., 
5, 1920, 224; Phytomonas angulata Bergey 



114 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



et al., Manual, 3rd ed., 1930, 267.) From 
L. angulatus, referring to the type of 
lesion produced on the tobacco leaf. 

Description taken from Clara (Cornell 
Agr. Exp. Sta. Mem. 159, 1934, 24). 

Rods : 0.75 to 1.5 by 1.5 to 3.0 microns. 
Motile by 1 to 6 polar flagella. Gram- 
negative. 

Gelatin: Liquefaction. 

Green fluorescent pigment produced. 

Beef -extract agar colonies : Dull white, 
circular, raised, smooth and glistening. 

Broth: Turbid in 36 hours and 
greenish. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Lipolytic action negative (Starr and 
Burkholder, Phytopath., 32, 1942, 601). 

Acid but not gas from glucose, galac- 
tose, fructose, mannose, arabinose, xy- 
lose, sucrose and mannitol. Alkaline reac- 
tion from salts of citric, malic, succinic and 
tartaric acid. Rhamnose, maltose, lac- 
tose , raffinose , glycerol , sali ci n , and aceti c , 
lactic and formic acids are not fermented. 

Starch not hydrolyzed. 

Slight growth in broth plus 5 to 6 per 
cent salt (Burkholder). 

Facultative anaerobe. 

Distinctive characters: Braun (Phyto- 
path., 27, 1937, 283) considers this species 
to be identical in culture with Pseudo- 
monas tabaci, but they differ in the type 
of disease they produce. 

Sources: Isolated by Fromme and 
Murray from small angular leaf spots on 
tobacco. 

Habitat : Causes the angular leaf spot 
of tobacco {Nicotiana tahacum). 

70. Pseudomonas aptata (Brown and 
Jamieson) Stevens. {Bacterium aptatum 
Brown and Jamieson, Jour. Agr. Res., 1, 
1913, 206; Phytomonas aptata Bergey et 
al., Manual, 1st ed., 1923, 184; Stevens, 
Plant Disease Fungi, New York, 1925, 
22.) From Latin aptatus adapted. 

Rods: 0.6 to 1.2 microns. Motile with 
bipolar flagella. Gram -negative. 



Green fluorescent pigment produced in 
culture. 

Gelatin : Liquefaction. 

Agar slants : Moderate growth along 
streak, filiform, whitish, glistening. 

Broth : Turbid : A pellicle formed. 

Milk : Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole not produced in 10 days. Slight 
amount found later. 

No HoS produced. 

Acid from glucose, galactose and 
sucrose. No acid from lactose, maltose 
and mannitol (Paine and Banfoot, Ann. 
Appl. Biol., 11, 1924,312). 

Starch not hydrolyzed. 

Slight growth in broth plus 7 per cent 
salt (Burkholder). 

Optimum temperature 27° to 28°C. 
Maximum 34° to 35°C. Minimum below 
1°C. 

Aerobic. 

Source : Isolated from diseased nas- 
turtium leaves from Virginia and diseased 
beet leaves from Utah. 

Habitat : Pathogenic on sugar beets, 
nasturtiums, and lettuce. 

71. Pseudomonas primulae (Ark and 
Gardner) Starr and Burkholder. (Phy- 
tomonas primulae Ark and Gardner, 
Phytopath., 26, 1936, 1053; Starr and 
Burkholder, Phytopath., 82, 1942, 601.) 
From L. primulus, first; M.L. Primula, 
a generic name. 

Rods: 0.51 to 0.73 by 1.0 to 3.16 
microns. Motile with a polar flagellum. 
Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Agar colonies : Round, convex, smooth, 
glistening, yellowish. 

Milk: Coagulated. 

Nitrites not produced from nitrates. 

Indole not produced. No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., S^, 1942,601). 

Acid but not gas from glucose, lactose, 
sucrose, maltose, galactose, arabinose. 



FAMILY PSEUDOMONADACEAE 



115 



glycerol, dulcitol and mannitol. Starch 
not hydrolyzed. 

Growth in broth plus 5 per cent salt. 

Optimum temperature 19° to 22°C. 
Maximum 34°C. Minimum 10°C. 

Optimum pH 6.8 to 7.0. Minimum 
4.5 to 5.0. 

Facultative anaerobe. 

Source : Isolated from leaf-spot of 
Primula polyantha. 

Habitat : Pathogenic on Primrd'a spp. 

72. Pseudomonas viridilivida (Brown) 
Holland. (Bacterium viridilividum 
Brown, Jour. Agr. Res., 4, 1915, 475; 
Holland, Jour. Bact., 5, 1920, 225; Phylo- 
monas viridilivida Bergey et al., ^Manual, 
1st ed., 1923, 187.) From Latin, viridis, 
green; lividus, blue. 

Rods : 1 .0 to 1 .25 by 1 .25 to 3.0 microns. 
Motile with 1 to 3 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin : Slow liquefaction. 

Beef agar colonies: Cream white, 
round, smooth, translucent, edges entire. 

Broth: Turbid, becomes lime green. 

Milk: Alkaline and clears. 

Nitrites not produced from nitrates. 

Indole is produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid from glucose and sucrose (Burk- 
holder). 

Grows well in 4.5 per cent salt. Grows 
in 7 per cent salt (Burkholder). 

Maximum temperature 34.5°C. Mini- 
mum 1.5°C. 

Aerobic. 

Source : Isolated from diseased lettuce 
from Louisiana. 

Habitat: Pathogenic on lettuce, Lac- 
tuca saliva. 

73. Pseudomonas delphinii (Smith) 
Stapp. (Bacillus delphini Smith, 
Science, 19, 1904, 417; Bacteriujii del- 
phinii Bryan, Jour. Agr. Res., 28, 1924, 
261 ; Stapp, in Sorauer, Handbuch der 
Pflanzenkrankheiten, 2, 5 Auf.. 1928. 106; 



Phytomonas delphinii Bergey et al., 
Manual, 3rd ed., 1930, 261.) From 
Latin, delphin, a dolphin; ^LL. Del- 
phinium, a generic name. 

Rods: 0.6 to 0.8 by 1.5 to 2.0 microns. 
Chains present. Motile with 1 to 6 polar 
flagella. Capsules. Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquified. 

Beef agar slants : Growth thin, smooth, 
shining, transparent, margins entire, 
crystals. Agar becomes dark brown. 

Broth : Turbid in 24 hours with delicate 
pellicle. 

Milk: Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Lipolytic action negative (Starr and 
Burkholder, Phytopath., 32, 1942, 601). 

Acid from glucose, galactose and 
fructose; slightly acid from sucrose. No 
acid from lactose, maltose, glycerol and 
mannitol. 

Starch : Hydrolysis feeble. 

Weak growth in broth plus 4 per cent 
salt. 

Optimum pH 6.7 to 7.1. pH range 5.6 
to 8.6. 

Optimum temperature 25°C. Maxi- 
mum 30°C. Minimum 1°C. or less. 

Source: Isolated from black spot of 
delphinium. 

Habitat: Pathogenic on delphinium 
causing a black spot in the leaves. 

74. Pseudomonas berberidis (Thorn- 
berry and Anderson) Stapp. (Phyto- 
monas berberidis Thornberry and Ander- 
son, Jour. Agr. Res., 43, 1931, 36; Stapp, 
Bot. Rev., 1, 1935, 407; Bacterium ber- 
beridis Burgwitz, Phytopathogenic Bac- 
teria, Leningrad, 1935, 153.) From ]\I.L. 
Berberis, barberry, generic name. 

Rods: 0.5 to 1.0 by 1.5 to 2.5 microns, 
occurring singly or in pairs. Motile with 
2 to 4 polar flagella. Capsules present. 

Gram-negative (Burkholder) ; not 
Gram-positive as stated in original de- 
scription. 



116 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Green fluorescent pigment produced in 
culture (Burkholder). 

Gelatin: Not liquefied. 

Glucose agar slants: Growth moder- 
ate, filiform at first, later beaded, raised, 
smooth, white. Butyrous in consistency. 

Milk: Becomes alkaline. No other 
change. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., S^, 1942,601). 

Acid from glucose, galactose, and 
sucrose. Maltose and rhamnose not 
utilized (Burkholder). 

No gas from carbohydrates. 

Starch not hydrolyzed. 

Optimum temperature 18°C. Maxi- 
mum 30°C . Minimum 7°C . 

Aerobic. 

Sources: Repeated isolations from 
leaves and twigs of barberry. 

Habitat: Pathogenic on barberry, Ber- 
beris thunbergerii and B. vulgaris. 

75. Pseudomonas coronafaciens (Elli- 
ott) Stapp. {Bacterium coronafaciens 
Elliott, Jour. Agr. Res., 19, 1920, 153; 
Phytomonas coronafaciens Bergey et al.. 
Manual, 1st ed., 1923, 180; Stapp, in 
Sorauer, Handbuch der Pflanzenkrank- 
heiten, 2, 5 Auf., 1928, 20.) From L. 
corona, halo ;/aciens, producing, referring 
to the lesions on oat blades. 

Probable synonyms: Elliott (Bact. 
Plant Pathogens, 1930, 122) lists as 
synonyms. Bacillus avenae (Russell, 
Johns Hopkins Univ. Thesis, 1892) and 
Bacillus avenae Manns and Pseudomonas 
avenae Manns (Ohio Agr. Exp. Sta. Bui. 
210, 1909, 133; Phytomonas avenae Ber- 
gey et al., Manual, 3rd ed., 1930, 263). 

Rods: 0.65 by 2.3 microns, occurring 
in chains. Motile with polar flagella. 
Capsules. Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin : Slow liquefaction. 

Nutrient agar colonies: White, becom- 



ing irregularly circular, flat with raised 
margins. 

Broth: Slight turbidity in 24 hours. 
Heavy pellicle formed. 

Milk: Alkaline. A soft curd formed 
followed by clearing. Curd sometimes 
absent. 

Nitrites not produced from nitrates. 

Indole not formed. 

No HjS formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 3^, 1942, 601). 

Acid but no gas from glucose and su- 
crose. Starch hydrolysis slight. 

Slight growth in broth plus 2 per cent 
salt. 

Optimum temperature 24° to 25°C. 
Maximum 3rC. Minimum 1°C. 

Source : Numerous isolations from 
blighted blades of oats. 

Habitat : Causes a halo spot on oats 
(Avena sativa). Artificial inoculations 
show barley {Hordeum vulgar e), rye 
{Secale cereale) and wheat (Triticum 
aeslivum) to be susceptible. 

75a. Pseudomonas coronafaciens var. 
atropurpiirea (Reddyand Godkin) Stapp. 
(Bacterium coronafaciens var. atropur- 
pureum Rcddy and Godkin, Phytopath., 
13, 1923, 81 ; Stapp, in Sorauer, Handbuch 
der Pflanzenkrankheiten, 2, 5 Auf., 1928, 
39; Phytomonas coronafaciens var. atro- 
purpurca Magrou, in Hauduroy et al., 
Diet. d. Bact. Path., Paris, 1937, 371.) 
From L. ater, black, dark; purpureus, 
purple, referring to the color of the lesion 
on brome grass. 

Distinctive characters : This variety 
differs from Pseudomonas coronafaciens 
in that it infects the brome-grass, Bromus 
inermis, where it produces a water soaked 
spot which is dark purple in color. 

Source : Numerous isolations from dis- 
eased brome-grass. 

Habitat : Pathogenic on Bromus 
inermis and Agropyron repens. Has been 
artificially inoculated on oats, Avena 
sativa. 



FAMILY PSEUDOMONADACEAE 



117 



76. Pseudomonas lachrymans (Smith 
and Bryan) Carsner. (Bacterium lachry- 
mans Smith and Bryan, Jour. Agr. Res., 
5, 1915, 466; Carsner, Jour. Agr. Res., 15, 
1918, 15; Bacillus lachrymans Holland, 
Jour. Bact., 5, 1920, 218; Phytomonas 
lachrymans Bergey et al., Manual, 1st ed., 
1923, 184.) From Latin, causing tears, 
probably referring to the opaque drops 
formed on the lesion caused by this 
pathogen. 

Synonym: Elliott (Man. Bact. Plant 
Pathogens, 1930, 147) lists the following 
as a synonym: Bacillus burgeri Potebnia, 
Khartov Prov. Agr. Exp. Sta., 1, 1915, 37. 
Description from Smith and Brj^an 
(loc. cil.) and Clara (Cornell Agr. Exp. 
Sta. Mem. 159, 1934, 26). 

Rods: 0.8 by 1 to 2 microns. Motile 
with 1 to 5 polar flagella. Capsules. 
Gram-negative. 

Green fluorescent pigment produced in 
culture. 
Gelatin: Liquefied. 

Beef-peptone agar colonies : Circular, 
smooth, glistening, transparent, whitish, 
entire margins. 

Broth: Turbid in 24 hours. White 
precipitate with crystals. 
Milk: Turns alkaline and clears. 
Nitrites not produced from nitrates. 
Indole reaction weak. 
No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but not gas from gUicose, fruc- 
tose, mannose, arabinose, .xylose, sucrose 
and mannitol. Alkaline reaction from 
salts of citric, malic and succinic acid. 
Maltose, rhamnose, lactose, raffinose, 
glycerol and salicin not fermented (Clara, 
loc. cit.). 

Starch partially digested. Not digested 
(Clara, loc. cit.). 

Growth in 3 per cent salt after 12 days. 
No growth in 4 per cent salt. 

Optimum temperature 25° to 27°C. 
Maximum 35°C. Minimum 1°C. 

Aerobic. Facultative anaerobe (Clara, 
loc. cit.). 



Source : Isolated from diseased cucum- 
ber leaves collected in New York, Wis- 
consin, Indiana and in Ontario, Canada. 

Habitat : Pathogenic on cucumber, 
Cucumis sativus, and related plants. 

77. Pseudomonas maculicola (McCul- 
loch) Stevens. {Bacterium maculicolum 
McCulloch, U. S. Dept. Agr., Bur. Plant 
Ind. Bui., 225, 1911, 14; Stevens, The 
Fungi which cause Plant Diseases, 1913, 
28; Phytomonas maculicola Bergey et al.. 
Manual, 1st ed., 1923, 189; Bacterium 
maccullochianum Burgwitz, Phytopatho- 
genic Bacteria, Leningrad, 1935, 77.) 
From L. maculus, spot; -cola, dweller. 

Rods: 0.9 by 1.5 to 3 microns. Fila- 
ments present. Motile with 1 to 5 polar 
flagella. Gram-negative. 

Green fluorescent pigment produced in 
culture. 
Gelatin: Liquefied. 
Beef -peptone agar colonies : Whitish, 
circular, shining, translucent, edges 
entire. 
Broth: Turbid. No ring or pellicle. 
Milk: Becomes alkaline and clears. 
Nitrites not produced from nitrates. 
Indole production feeble. 
No H-S formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid from glucose, galactose, xylose, 
sucrose, glycerol, and mannitol. Alka- 
line reaction from salts of citric, malic, 
malonic, and succinic acid. Salicin, 
maltose, and salts of hippuric and tartaric 
not utilized (Burkholder). 

Slight growth in broth plus 4 per cent 
salt (Erw. Smith, Bact. Plant Diseases, 
1920, 306). 
Aerobic. 

Optimum temperature 24° to 25°C. 
Maximum 29°C. Minimum 0°C. 

Source : Isolated from diseased cauli- 
flower leaves from Virginia. 

Habitat : Pathogenic on cauliflower and 
cabbage. 

Note: Bacterium maculicola var.japo- 
nicum Takimoto, Bui. Sci. Fak. Terkult 



118 



MANUAL or DETERMINATIVE BACTERIOLOGY 



Kjusu Imp. Univ., 4, 1931, 545 has not 
been seen. 

78. Pseudomonas marginata (McCul- 
loch) Stapp. {Bacterium viarginatum 
McCulloch, Science, 5If, 1921, 115; Jour. 
Agr. Res., 29, 1924, 174; Phytomon'as 
marginata Bergey et al.. Manual, 1st ed., 
1923, 188; Stapp, in Sorauer, Handbuch 
der Pflanzenkrankheiten, 2, 5 Auf., 1928, 
56.) From L. marginatus, having a 
border, probably refers to the definite 
margin of the colony. 

Rods: 0.5 to 0.6 by 0.8 to 1.8 microns. 
Motile with 1 to 4 bipolar flagella. Cap- 
sules. Gram-negative. 

Green fluorescent pigment produced in 
Uschinsky's and Fermi's solutions. 

Gelatin: Liquefied. 

Agar colonies : White, circular, smooth, 
translucent, viscid, with definite margins 
at first thin but later thick and con- 
toured. Surface wrinkled. 

Milk: At first slightly acid, then alka- 
line. Casein digested. 

Nitrites not produced from nitrates. 

Indole production slight. 

Hydrogen sulfide production slight. 

Lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but not gas from glucose, lactose, 
sucrose and glycerol. 

Starch hydrolysis feeble. 

Growth in 3.5 per cent salt. No growth 
in 4 per cent salt. pH range, 4.6 to 9.1. 

Optimum temperature 30° to 32°C. 
Maximum 40°C. Minimum 8° to 9°C. 

Source : Repeatedly isolated from 
diseased gladiolus. 

Habitat: Pathogenic on Gladiolus spp. 
and Iris spp. 

79. Pseudomonas medicaginis Sackett. 
(Sackett, Science, 31, 1910, 553; also 
Colorado Agr. Exp. Sta., Bull. 158, 1910, 
11 ; Bacillus medicaginis Holland, Jour. 
Bact., 5, 1920, 219; Phytomonas medi- 
caginis Bergey et al., Manual, 1st ed., 
1923, 179; Bacterium medicaginis Elliott, 
Bact. Plant Path., 1930, 162.) From L. 



medica, ancient Media; M.L. Medicago, a 
generic name. 

Rods : 0.7 by 1 .2 microns. Motile with 
1 to 4 flagella. Filaments present. 
Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Not liquefied. 

Nutrient agar colonies: Growth in 24 
hours whitish, glistening. 

Broth: Turbid in 24 hours. Pellicle 
formed. Viscid sediment. 

Milk: Becomes alkaline. No change. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Starch not hydrolyzed. 

No gas from carbohydrates. Acid from 
sucrose. 

Slight growth in broth plus 3.75 per cent 
salt. 

Optimum temperature 28° to 30°. 
Maximum 37.5°C. 

Aerobic. 

Source : Isolated from brown lesions on 
leaves and stems of alfalfa. 

Habitat : Pathogenic on alfalfa, Medi- 
cago sp. 

79a. Pseudomonas phaseolicola (Burk- 
holder) Dowson. {Phytomonas medi- 
caginis var. phaseolicola Burkholder, 
Phytopath., 16, 1926, 915; Bacterium 
medicaginis var. phaseolicola Link and 
Hull, Bot. Gaz., 83, 1927, 413; Pseudo- 
monas medicaginis var. phaseolicola 
Stapp and Kotte, Nachrichtenb. f. d. 
Deutschen Pflanzenschutzdienst, 9, 1929, 
35; Dowson, Brit. Mycol. Soc. Trans., 
26, 1943, 10.) From L. phaseolus, bean; 
M.L. Phaseolus, a generic name;-coZa, 
dweller. 

Synonym : Bacterium puerariae 
Hedges. (Phytopath., 17, 1927, 48 and 
20, 1930, 140; Phytomonas puerariae 
Bergey et al.. Manual, 3rd ed., 1930, 267.) 

Description from Burkholder and 
Zaleski (Phytopath., 22, 1932, 85). 



FAMILY PSEUDOMONADACEAE 



119 



Rods : 1 by 2 microns, sometimes 
slightly curved, filaments present. 
Motile with polar flagellum. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin stab: Slow liquefaction. 

Beef extract agar: Whitish, circular 
colonies, 2 mm. in diameter. Edges 
entire. 

Broth: Turbid. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not formed. 

Not lipolytic (Starr and Burkholder, 
Photopath., 35, 1942, 601). 

Acid but no gas from glucose, fructose, 
mannose, arabinose, xjdose, sucrose and 
glycerol. No acid from rhamnose, lac- 
tose, maltose, mannitol and salicin. 
Alkali from salts of citric and malic acids, 
but not from acetic, formic, lactic or 
tartaric acids. Starch and cellulose not 
hydrolyzed. 

Slight growth in broth plus 4 per cent 
salt. 

Optimum temperature 20"" to 23°C. 
Maximum 33°C. Minimum 2.5°C. 
(Hedges, loc. cit.). 

Optimum pH 6.7 to 7.3. Maximum S.8 
to 9.2. Minimum 5.0 to 5.3. (Kotte, 
Phyt. Zeitsch., 2, 1930, 453.) 

Microaerophilic. 

Source : Isolated from leaves, pod and 
stem of beans showing halo blight. 

Habitat: Pathogenic on beans (Phaseo- 
lus vulgaris), the kudzu vine (Pueraria 
hirsula) and related plants. 

80. Pseudomonas pisi Sackett. (Sack- 
ett, Colorado Agr. E.xp. Sta., Bull. 218, 
1916, 19; Bacterium -pisi Erw. Smith, 
An Introduction to Bacterial Diseases of 
Plants, 1920, 474; Phytomonas pisi Ber- 
gey et al.. Manual, 1st ed., 1923, 181.) 
From Gr. pisum, the pea; M.L. Pisum, 
a generic name. 

Rods : 0.68 to 2.26 microns. Motile 
with a polar flagellum. Gram -negative. 



Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Agar slants : Moderate growth in 24 
hours, filiform, glistening, grayish-white. 

Broth: Turbid with a scum in 5 days. 

Milk: Alkaline, soft curd, clears. 

Nitrites not produced from nitrates. 

Indole not produced! 

No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but notgas from glucose, galactose 
and sucrose. 

Starch not hydrolyzed. 

Optimum temperature 27" to 28°C. 
Maximum 37.5°C. Minimum 7°C. 

Aerobic. 

Source : Ten cultures isolated from 5 
collections of diseased peas showing water 
soaked lesions on stems and petioles. 

Habitat: Pathogenic on garden peas, 
Pisum sativum and field peas, P. sativum 
var. arvense. 

81. Pseudomonas syringae van Hall. 
(Kennis der Bakter. Pfianzenziekte, 
Inaug. Diss., Amsterdam, 1902, 191; 
Bacterium syringae Erw. Smith, Bacteria 
in Relation to Plant Diseases, 1, 1905, 68; 
Phytomonas syringae Bergey et al., 
Manual, 3rd ed., 1930, 257.) From 
Latin, syringa, a nymph that was 
changed into a reed; M.L. Syringa, a 
generic name. 

Synonyms: Bryan (Jour. Agr. Res., 38, 
1928, 225) lists Bacterium citriputeale C. 
O. Smith, Phytopath., 3, 1913, 69, and 
Bacterium citrarefaciens Lee, Jour. Agr. 
Res., 9, 1917, 1 {Pseudomonas citrare- 
faciens Stapp, in Sorauer, Handb. d. 
Pflanzenkrankheiten, 2, 5 Aufl., 1928, 
190). Clara (Cornell Agr. Exp. Sta. 
Mem. 159, 1934, 29) lists Bacterium vignae 
{Pseudomonas vignae) Gardner and Ken- 
drick, Science, 57, 1923, 275 {Phytomonas 
vignae Bergey et al., Manual, 1st ed., 
1923, 188), Pseudomonas viridifaciens 
Tisdale and Williamson, Jour. Agr. Res., 
25, 1923, 141 .{Bacterium viridifaciens 
Tisdale and Williams, ibid. ; Phytomonas 



120 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



viridifaciens Bergey et al., Manual, 2nd 
ed., 1925, 208), and Phylomonas vignae 
var. leguminophila Burkholder, Cornell 
Agr. Exp. Sta. Mem. 127, 1930, 51. Wil- 
son (Phytopath., 30, 1940, 27) lists 
Phytomonas cerasi (Griffin) Bergey et al. 
(Pseudomonas cerasus Griffin, Science, 
34, 1911, 615; Bacillus cerasus Holland, 
Jour. Bact., 5, 1920, 217; Bergey et al., 
Manual, 3rd ed., 1930, 262; Bacterium 
cerasi Elliott, Bact. Plant Pathogens, 
1930, 109.) This would include, there- 
fore the following synonyms which 
have been listed for Phytomonas cerasi. 
Clara (Cornell Agr. Exp. Sta. Mem. 
159, 1934, 25) lists Bacterium trifoliorum 
Jones et al. (Jour. Agr. Res., 25, 1923, 
471; Phytomonas trifoliorum Burk- 
holder, Phytopath., 16, 1926, 922; 
Pseudomonas trifoliorum Stapp, in Sor- 
auer, Handb. d. Pflanzenkrankheiten, 2, 
5 Aufl., 1928, 177) and Bacterium hold 
Kendrick (Phytopath., 16, 1926, 236; 
Pseudomonas hold Kendrick, ibid. ; 
Phytomonas hold Bergey et al.. Manual, 
3rd ed., 1930, 258). Wilson (Hilgardia, 
10, 1936, 213) lists Pseudomonas prunicola 
Wormald (Ann. Appl. Biol., 17, 1930, 
725), Pseudomonas cerasi var. prunicola 
Wilson (Hilgardia, S, 1933,83), Bacterium 
citripntcale C. 0. Smith (Phytopath., 4, 
1913, 69; Pseudomonas citriputealis 
Stapp, in Sorauer, Handb. d. Pflanzen- 
krankheiten, 2, 5 Aufl., 1928, 190; Phyto- 
monas citriputealis Bergey et al., Manual, 
3rd ed., 1930, 278) and Pseudomonas 
utiformica Clara, Science, 75, 1932, HI 
(Phytomonas utiformica Clara, Cornell 
Agr. Exp. Sta. Mem. 159, 1934, 29; Bac- 
terium utiformica Burgwitz, Phytopatho- 
genic Bacteria, Leningrad, 1935, 444). A 
probable synonym is Phytomonas 
spongiosa (Aderhold and Ruhland) Ma- 
grou {Bacillus spongiosus Aderhold 
and Ruhland, Cent. f. Bakt., II Abt., 15, 
1905, 376; Pseudomonas spongiosa Braun, 
Die Landwirtschaft, 41, 42, 1927, 2 pp.; 
Bacterium spongiosum Elliott, Man. 
Bact. Plant Pathogens, 1930, 214; Ma- 
grou, in Hauduroy et al.. Diet. d. Bact. 
Path., Paris, 1937, 414). C. O. Smith 



(Phytopath., S3, 1943, 82) lists the follow- 
ing as a synonym : Pseudomonas hibisci 
(Nakata and Takimoto) Stapp (Bacteri- 
um hibisci Nakata and Takimoto, Ann. 
Phytopath. Soc. Japan, 1, 5, 1923, 18; 
Stapp, in Sorauer, Handbuch der Pflan- 
zenkrankheiten, 5 Aufl., 2, 1928, 203; 
Phytomonas hibisci Bergey et al., Manuals 
3rd ed., 1930, 264). 

Description from Clara (loc. cit.). 

Rods: 0.75 to 1.5 by 1.5 to 3.0 microns. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Beef -extract agar colonies : Circular, 
grayish-white with bluish tinge. Surface 
smooth. Edges entire or irregular. 

Broth: Turbid in 36 hours. No pelli- 
cle. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not produced. 

No HoS produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Slight growth in broth plus 4 per 
cent salt. 

Acid but not gas from glucose, galac- 
tose, mannose, arabinose, xylose, su- 
crose, mannitol and glycerol. Alkaline 
reaction from salts of citric, malic, suc- 
cinic and lactic acid. Rhamnose, mal- 
tose, lactose, raffinose, salicin, and acetic, 
formic and tartaric acid not fermented. 

Starch not hydrolyzed. 

Facultative anaerobe. 

Source : Van Hall originally isolated 
the pathogen from lilac. 

Habitat : Pathogenic on lilac, citrus, 
cow peas, beans, lemons, cherries and 
many unrelated plants. 

81a. Orsini reports the following as a 
variety — Bacterium syringae var. capsici 
Orsini. (Intern. Bull. Plant Prot., 33, 
1942, 33.) Pathogenic on the pepper 
plant (Capsicum). 

82. Pseudomonas atrofaciens (McCul- 
loch) Stevens. (Bacterium atrofaciens 



FAMILY PSEUDOMONADACEAE 



121 



McCulloch, Jour. Agr. Res., 18, 1920, 
549 ; Phytomonas atrofaciens Bergey et 
al., Manual, 1st ed., 1923, 185; Stevens, 
Plant Disease Fungi, New York, 1925, 22.) 
From Latin, ater, black; faciens, making, 
referring to the color of the lesion on 
wheat. 

Rods: 0.6 by 1 to 2.7 microns. Long 
chains formed in culture. Capsules 
present. Motile with 1 to 4 polar or bi- 
polar flagella. Gram -negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: Liquefied. 

Beef-peptone-agar colonies: Circular, 
shining, translucent, white. 

Broth: Growth never heavy, slight 
rim, and a delicate pellicle. 

Milk: Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole: Slight production. 

Hj^drogen sulfide: Slight production. 

Acid and no gas from glucose, galac- 
tose and sucrose. 

Starch is slightly hytlrolyzed. 

Optimum temperature 25° to 28°C. 
Maximum 36° to 37°C and minimum be- 
low 2°C. 

Aerobic. 

Sources : Isolated from diseased wheat 
grains collected throughout United States 
and Canada. 

Habitat: Causes a basal glume-rot of 
wheat. 

83. Pseudomonas cumini (Kovacev- 
ski) Dowson. {Phytomonas cumini Kov- 
acevski. Bull. Soc. Bot. Bulgarie, 7, 1936, 
27; Dowson, Trans. Brit. Mycol. Soc, 
26, 1943, 10.) From Greek cuminum, 
cumin; M.L. Cuminum, a generic name. 

Rods : 0.5 to 0.7 by 1 to 3 microns, oc- 
curring in chains and filaments. Motile 
with 1 to 3 polar flagella. Gram-negative. 

Green fluorescent pigment formed in 
culture. 

Gelatin: Rapidly liquefied. 

Potato agar colonies: Grayish-white, 
circular, glistening, smooth, butyrous. 

Broth: Moderate turbidity. Pseudo- 
zoogloea. 



Milk: Xot coagulated. Casein pep- 
tonized. 

Nitrites not produced from nitrates. 

Indole not formed. 

No HaS formed. 

Acid but not gas from glucose and 
sucrose. No acid from lactose or glyc- 
erol. Starch not hydrolyzed. 
■' Temperature range 5°C to 31°C. 

Aerobic. 

Source : Isolated from blighted cumin 
(Cuminum). 

Habitat : Pathogenic on cumin and dill. 

84. Pseudomonas desaiana (Burk- 
holder) comb. nov. (B. pyocyancus sac- 
charum Desai, Ind. Jour. Agr. Sci., 5, 
1935, 391 ; Phytomomas desaiana Burk- 
holder, in Bergey et al., Manual, 5th ed., 
1939, 174.) Named for Desai who first 
isolated the species. 

Rods: 0.6 to 1.2 by 1.2 to 2.2 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: Liquefaction. 

Agar colonies: Grayish-blue. Raised. 

Broth: Light clouding. Pellicle. 

Milk: Peptonized without coagulation. 

Nitrites not produced from nitrates. 

Indole not formed. 

Glucose, sucrose, lactose and glycerol 
fermented without gas. 

Starch: Hydrolysis present. 

Optimum temperature 30°C. 

Aerobic. 

Source : Isolated from stinking rot of 
sugar cane in India and associated with a 
white non-pathogenic bacterium. 

Habitat : Pathogenic on sugar cane, 
Saccharu tn officinar urn . 

85. Pseudomonas erodii Lewis. (Phy- 
topath., 4, 1914, 231; Bacterium erodii 
Lewis, ibid.; Phytomonas erodii Bergey 
et al., ^'lanual, 3rd ed., 1930, 256.) From 
Greek, erodius, heron; ]\I.L. Erodium, 
a generic name. 

Rods: 0.6 to 0.8 by 1.2 to 1.8 microns. 



122 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Motile with 1 to 3 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: Liquefaction. 

Agar stroke : Heavy, smooth, cream - 
colored growth in 24 hours. 

Broth: Dense clouding in 24 hours. 

Milk : Turns alkaline and clears, litmus 
reduced. 

Nitrites not produced from nitrates. 

Indole produced in 14 days. 

No H2S produced. 

Acid but not gas from glucose, sucrose, 
lactose and glycerol. 

Temperature: No growth at 35°C. 

Aerobic, obligate. 

Source: Isolations from Erodium tex- 
anum and 4 varieties of Pelargonium. 

Habitat : Causes a leaf spot of Erodium 
iexanum and Pelargonium spp. 

86. Pseudomonas apii Jagger. (Jagger, 
Jour. Agr. Res., £1, 1921, 186; Phyto- 
monas apii Bergey et al.. Manual, 1st ed., 
1923, 184; Pseudomonas jaggeri Stapp, in 
Sorauer, Handbuch der Pflanzenkrank- 
heiten, 2, 5 Aufl., 1928, 210; Bacterium 
jaggeri Elliott, Bacterial Plant Patho- 
gens, 1930, 142; Phytomonas jaggeri 
Magrou, in Handuroy et al., Diet. d. 
Bact. Path., Paris, 1937, 371.) From 
Latin, apium, parsley, M.L. Apium, a 
generic name. 

Description from Clara (Cornell Agr. 
Exp. Sta. Mem. 159, 1934, 24). 

Rods: 0.75 to 1.5 by 1.5 to 3.0 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Green fluorescent pigment produced 
in various media. 

Gelatin: Liquefaction. 

Beef -extract agar colonies: Circular, 
glistening, smooth, edges entire. Gray- 
ish-white with bluish tinge. 

Broth: Turbid in 36 hours. PeUicle 
formed. 

Milk: Becomes alkaline. No curd. 

Nitrites not produced from nitrates. 

Indole not formed. 



No H2S formed. 

Acid but not gas from glucose, galac- 
tose, fructose, mannose, arabinose, 
xylose, sucrose, mannitol and glycerol. 
Alkaline reaction from salts of acetic, 
citric, malic and succinic acids. Rham- 
nose, maltose, lactose, raffinose, salicin, 
and formic, lactic and tartaric acid are 
not utilized. 

Starch not hydrolyzed. 

Facultative anaerobe. 

Distinctive characters : Pathogenicity 
appears limited to celery. 

Source : Jagger isolated this repeatedly 
from diseased celery leaves. 

Habitat: Pathogenic on celery, Apium 
graveolens. 

87. Pseudomonas matthiolae (Briosi 
and Pavarino) Dowson. {Bacterium mat- 
thiolae Briosi and Pavarino, Atti della 
Reale Accad. dei Lincei Rend., 21 , 1912, 
216; Phytomonas matthiolae, Bergey et 
al.. Manual, 3rd ed., 1930, 266; Mushin, 
Proc. Roy. Soc. Victoria, 53, 1941, 201; 
Dowson, Trans. Brit. Mycol. Soc, 26, 
1943, 10.) From M.L. Matthiola, a 
generic name. 

Rods: 0.4 to 0.6 by 2 to 4 microns. 
Gram-positive. Gram-negative (Mush- 
in, loc. cit.). 

Green fluorescent pigment produced 
in culture. 

Gelatin : Liquefied. 

Beef agar colonies : White, circular 
colonies, slightly elevated, margins 
smooth. 

Broth: Slightly turbid. Becomes pale 
green. 

Milk: Coagulation with acid reacfon. 

Nitrites produced from nitrates 
(Mushin). 

Hydrogen sulfide not formed. 

Acid from glucose, galactose, fructose, 
mannose, rhamnose, glycerol, mannitol, 
acetic acid, citric acid, formic acid, lactic 
acid, malic acid, and succinic acid. 
Feeble acid in maltose. No acid, no gas 
in lactose, sucrose, raffinose, starch, 
salicin, and tartaric acid (Mushin). 

Optimimi temperature 20 to 24°C. 



FAMILY PSEUDOMONADACEAE 



123 



Maximum temperature 38.5 C. Mini- 
mum below 0°C. (Mushin). 

Limits of growth in broth are pH 4.4 
to pH 9.5 (Mushin). 

Aerobic. 

Source: Isolated from vascular and 
parenchymatic disease of stocks, Mat- 
thiola incana var. annua. 

Habitat: Pathogenic on stocks. 

Note: Burkholder (Phytopath., 28, 
1938, 936) and Santarelli (Rev. di Pat. 
Veg., 29, 1939, 364) consider this species 
a synonym of Pseudomonas syringae. 
Adam and Pugsley (Jour. Dept. Agric. 
Victoria, 32, 1934, 306) give a descrip- 
tion of a green fluorescent pathogen on 
stocks which is similar to Pseudomonas 
syringae. Mushin {loc. cit.) considers 
Pseudomonas matthiolae to be a distinct 
species. 

88. Pseudomonas mors-prunorum 
Wormald. (Jour. Pom. and Hort. Sci., 
9, 1931,251; Phytomonas mors-prunorum 
Wormald, Trans. Brit. Mycol. Soc, 17, 
1932, 169; Bacterium mors-prunorum, 
ibid.) From L. mors, death; prunus, 
plums. 

Rods: Motile with a polar flagellum. 
Gram-positive (1931). Gram-negative 
(1932). 

Note : Possibly a green fluorescent or- 
ganism since it produces a faint yellow 
color in Uschinsky's solution. 

Gelatin: Liquefaction. 

Agar colonies : White. 

Broth plus 5 per cent sucrose: White 
and cloudy. 

Nitrites not produced from nitrates. 

Acid but not gas from glucose, lactose, 
sucrose and glycerol. 

Starch not hydrolyzed. 

Strict aerobe. 

Distinctive characters : Differs from 
Pseudomonas prunicola {Pseudomonas 
syringae) in that it produces a white 
cloudy growth in broth plus 5 per cent 
sucrose; a rapid acid production in nu- 
trient agar plus 5 per cent sucrose, and a 
faint yellow or no color in Uschinsky's 
solution. 



Source : Isolated from cankers on plum 
trees in England. 
Habitat : Pathogenic on Prunus spp. 

89. Pseudomonas rimaefaciens Koning. 

(Chron. Bot., 4, 1938, 11 ; Meded. Phytop. 
Labor, Willie Comm. Scholt., 14, 1938, 
24.) From L. rima, fissure; Jaciens, pro- 
ducing. 

Rods: 0.6 to 2.4 microns in length. 
Motile with 1 to 3 polar flagella. Gram- 
negative. 

Yellow -green fluorescent water-soluble 
pigment produced in culture. 

Gelatin : Liquified. 

Agar colonies: Round, convex, smooth, 
somewhat granular with hyaline edge. 

Broth : Turbid. Surface growth with 
a sediment in a few days. 

Milk: Alkaline and clears. 

Nitrites not produced from nitrates. 
Peptone, asparagin, urea, gelatin, ni- 
trates and ammonia salts are sources of 
nitrogen. 

Hydrogen sulfide not produced. 

Indole production slight. 

Growth with the following carbon 
sources plus NO3, glucose, sucrose, 
glycerol, succinates, malates, citrates 
and oxalates. Less growth with manni- 
tol, fructose, galactose, lactose, salicy- 
late. Acid is produced from the sugars. 
No growth with dextrin, inulin, maltose, 
lactose, rhamnose, salicin, tartrates, 
acetates, formates. 

Starch not hj^drolyzed. 

Aerobic. 

Optimum temperature 25°C. Maxi- 
mum about 37°C. Very slow growth at 
14°C. Thermal death point 42° to 48°C. 

Source: Strains of the pathogen iso- 
ated from poplar cankers in France and 
in the Netherlands. 

Habitat : Pathogenic on Populus bra- 
bantica, P. trichocarpa and P. candicans. 

This may be Pseudomonas syringae 
since the characters are the same and 
both organisms can infect Impatiens sp. 
Pseudomonas syringae infects poplars 
(Elliott, Bacterial Plant Pathogens, 
1930, 218). 



124 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



90. Pseudomonas papulans Rose. 
(Rose, Phytopath., 7, 1917, 198; Phyto- 
monas papulans Bergey et al.. Manual, 
3rd ed., 1930, 267; Bacierivrn papulans 
Elliott, Bacterial Plant Pathogens, 1930, 
175; Phytonionas syringae var. papulans 
Smith, Jour. Agr. Res., 68, 1944, 294.) 
From L. papulans, forming blisters. 

Rods : 0.6 by 0.9 to 2.3 microns. Motile 
with 1 to 6 polar flagella. Gram-nega- 
tive. 

Green fluorescent pigment produced in 
culture. 

Gelatin : Liquefied. 

Broth : Moderate turbidity in 24 hours. 

Milk : Alkaline and at times a soft 
coagulum. 

Nitrites not produced from nitrates. 

Indole: May or may not be produced. 

Acid but not gas formed from glucose 
and sucrose. 

Optimum temperature 25° to 28°C. 
Maximum 37°C. 

Source : Twenty-five cultures isolated 
from blisters on apples and from rough 
bark. 

Habitat: Pathogenic on apple trees. 

91. Pseudomonas pseudozoogloeae 

(Honing) Stapp. (Baclcrium pseudo- 
zoogloeae Honing, Bull, van Het. Deli 
Proefstation, Medan, 1, 1914, 7; Stapp, 
in Sorauer, Handbuch der Pflanzen- 
krankheiten, 2, 5 Auf., 1928, 274; Phyto- 
rnonas pseudozoogloeae Bergey et al., 
3rd ed., 1930, 261.) From Gr., pseudo, 
false; M. L. zoogloea, zooglea. 

Rods: 0.7 to 1.5 by 0.9 to 2.5 microns. 
Chains. Motile with 1 or 2 polar flagella. 
Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Agar colonies: Round, flat, yellow- 
gray. 

Broth : Moderate turbidity with pseu- 
dozoogloea in the pellicle. 

Milk : Coagulation. No clearing. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced. 



Acid but not gas from glucose, lactose, 
maltose, sucrose and mannitol. 

Facultative anaerobe. 

Source : Isolated from the black rust 
of tobacco. 

Habitat : Pathogenic on tobacco, 
Nicotiana tabacum. 

92. Pseudomonas tabaci (Wolf and 
Foster) Stevens. {Bacterium tabacum 
Wolf and Foster, Science, ^6, 1917, 362; 
also Jour. Agr. Res., 12, 1918, 449; Phyto- 
monas tabaci Bergey et al., Manual, 1st 
ed., 1923, 185; Stevens, Plant Disease 
Fungi, New York, 1925, 34.) From 
Nicotiana tabacum, tobacco. 

Rods : 1.2 by 3.3 microns. Motile with 
a polar flagellum. Gram-negative. 

Gelatin : Liquefaction. 

Potato agar colonies: Grayish-white, 
circular, raised, wet-shining, smooth. 

Milk: Alkaline; clears. 

Nitrites not produced from nitrates. 

Indole not formed. 

Acid from glucose, galactose, fructose, 
1-arabinose, xylose, sucrose, pectin, man- 
nitol and glycerol (Braun, Phytopath., 
27, 1937,289). 

Ammonium sulfate, potassium nitrate, 
cystine, glutamic acid, glycine, succin- 
imide, oxamide, acetamide, and urea can 
be used as nitrogen source (Braun). 

Starch not hydrolyzed. Aerobic. 

Distinctive character: Braun {loc. cit.) 
states that Pseudomonas tabaci and Pseu- 
domonas angulata are identical in culture. 

Source : Isolated from wildfire lesions 
on tobacco leaves in North Carolina. 

Habitat: Pathogenic on tobacco, Nico- 
tiana tabacum. 

93. Pseudomonas lapsa (Ark) Burk- 
holder. (Phytomonas lapsa Ark, Phyto- 
path., 30, 1940, 1; Burkholder, ibid., 32, 
1942, 601.) From Latin, lapsus, falling, 
referring to a symptom of the disease. 

Rods: 0.56 by 1.55 microns. Motile 
with 1 to 4 polar flagella. 

Produces fluorescence in Uschinsky's, 
Fermi's, and Cohn's solutions. 



FAMILY PSEUDOMONADACEAE 



125 



Gelatin: Liquefied (Burkholder). 

Acid but no gas is produced from 
glucose, sucrose, maltose, lactose, glyc- 
erine, arabinose, xylose, galactose, raf- 
finose and mannitol. 

Slight growth in broth plus 5 per cent 
salt (Burkholder). 

Source : Isolated from stalk rot of field 
corn in California; also from Diabrotica 
beetles. 

Habitat : Pathogenic on corn and sugar 
cane. 

Note : Like Pseudomonas desiana. 

94. Pseudomonas bowlesiae (Lewis 
and \Yatson) Dowson. (Phytomonas 
boidesii Lewis and Watson, Phytopath.. 
17, 1927, 511; Bacterimn bowlesii Elliott, 
Bacterial Plant Pathogens, 1930, 9G; 
Dowson, Trans. Brit. IMycol. Soc, 26, 
1943, 9.) From 'SI. L. Boidcsia. a generic 
name. 

Rods: 0.5 to 0.7 by 1.2 to 1.6 microns, 
occurring singl}^ in pairs or in short 
chains. Motile with bipolar flagella. 
Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin : Liquefied. 

Agar slants: Yellowish, moist, glisten- 
ing and viscid. 

Broth: Uniform turbidity- throughout. 
Heavy viscous sediment in old cultures. 

Milk: Alkaline; coagulation, with a 
slow peptonization. 

Nitrites are produced from nitrates. 

Indole is produced. 

Hydrogen sulfide is produced. 

Acid from glucose, maltose and xylose. 
No acid from sucrose. 

Optimum temperature 27"C. Maxi- 
mum 37°C. Minimum — l^C. 

Optimum pH 7.2. pH range 4.5 to 8.6. 

Aerobic. 

Source: Isolated from diseased, water 
soaked spots of bowlesia. 

Habitat : Pathogenic on Bowlesia sep- 
tentrionalis. 

95. Pseudomonas intybi (Swingle) 
Stapp. {Phytomonas intybi Swingle, 



Phytopath., 15, 1925, 730; Stapp, in Sor- 
aurer, Handbuch der Pflanzenkrank- 
heiten, 2, 5 Auf., 1928, 291; Bacterium 
intybi Elliott, Bacterial Plant Pathogens, 
1930, 142.) From Latin, intibus, endive. 

Description from Stapp, Cent. f. Bakt., 
II Abt., 9/, 1935, 232. 

Rods: 0.4 to 0.5 by 1.4 to 2.8 microns. 
Motile with one to several polar flagella. 
Gram-negative. 

Green fluorescent pigment formed in 
culture. 

Gelatin: Liquefaction. 

Agar colonies : White, glistening, trans- 
parent. 

Broth: Turbid with fragile pellicle, 
and good sediment. 

Milk: Coagulated. Casein not pep- 
tonized. 

Nitrites are produced from nitrates 
with the formation of gas. 

Indole not formed. 

Acid but not gas from arabinose, xylose 
and glucose. No acid from sucrose. 

Optimum temperature 23° to 28°C. 
Maximum 40° to 42°C. Minimum 0°C. 

Distinctive character: Differs from 
Psexidomonas cichorii in that it liquefies 
gelatin and produces nitrites from ni- 
trates. 

Source : Isolated from French endive, 
Cichorium inlybus by Swingle, from C. 
endiva and lettuce, Lactuca saliva by 
Stapp. 

Habitat : Pathogenic on endive and let- 
tuce, causing a rot. 

96. Pseudomonas marginalis (Brown) 
Stevens. (Bacterium marginale Brown, 
Jour. Agr. Res., 13, 1918, SSQ;Phylo}nonas 
marginalis Bergey et al.. Manual, 1st ed., 
1923, 182; Stevens, Plant Disease Fungi, 
New York, 1925, 30.) From Latin, margo 
{marginis), edge, margin; M.L. viar- 
ginalis, on the margin, a character of the 
disease. 

Description from Brown (loc. cit.) and 
Clara (Cornell Agr. Exp. Sta. Mem. 159, 
1934, 27). 

Rods : Motile with 1 to 3 polar flagella. 
Gram-negative. 



126 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Agar colonies: Cream-colored to yel- 
lowish. 

Brotla: Turbid, with pellicle. 

Milk: Alkaline. Soft curd at times. 

Nitrites are produced from nitrates. 
Not produced (Clara, loc. cit.). 

Indole not produced. 

No H2S produced. 

Acid but not gas from glucose, galac- 
tose, fructose, mannose, arabinose, xylose, 
rhamnose, mannitol and glycerol. Al- 
kali from salts of acetic, citric, malic, 
formic, lactic, succinic and tartaric acid. 
Sucrose, maltose, lactose, raffinose and 
salicin not fermented (Clara, loc. cit.). 

Starch hydrolysis feeble. None (Clara, 
loc. cit.). 

Optimum temperature 25° to 26°C. 
Maximum 38°C. Minimum 0°C. 

Aerobic. 

Source : Isolated from marginal lesion 
on lettuce from Kansas. 

Habitat : Pathogenic on lettuce and 
related plants. 

97. Pseudomonas setariae (Okabe) 
comb. nov. (Bacterium setariae Okabe, 
Jour. Soc. Trop. Agr. Formosa, 6, 1934, 
63; Phytomonas setariae Burkholder, in 
Bergey, Manual, 5th ed., 1939, -183.) 
From L. seia, bristle; -arius, like; M. L. 
Setaria, a generic name. 

Rods: 0.4 to 0.8 by 1.8 to 4.4 microns. 
Motile with a polar, seldom bipolar, flagel- 
lum. Gram-negative. 

Yellowish water-soluble pigment pro- 
duced in culture. 

Gelatin: Slow liquefaction. 

Beef -extract agar colonies : Circular, 
white, opalescent, smooth, glistening. 

Broth: Turbid after 18 hours. Pel- 
licle. 

Milk: Alkaline; clears. 

Nitrites are produced from nitrates. 

Indole is produced. 

No H2S produced. 

Acid but not gas from glucose, galac- 



tose and glycerol. No acid from lactose, 
maltose or sucrose. 

Starch: Feeble hydrolysis. 

Grows in 3 per cent salt. 

Optimum temperature 31° to 34°C. 
Maximum 42°C. 

Aerobic. 

Source : Isolated from brown stripe of 
Italian millet. 

Habitat: Pathogenic on Italian millet, 
Setaria italica. 

98. Pseudomonas polycolor Clara. 
(Clara, Phytopath., 20, 1930, 704; Phyto- 
monas polycolor Clara, ibid., Bacterium 
polycolor Burgwitz, Phytopathogenic 
Bacteria, Leningrad, 1935, 148.) From 
Gr. poly, many; L. color, color. 

Note: Delacroix (Comp. rend. Acad. 
Sci., Paris, 137, 1903, 454) describes 
Bacilhis aerogenosus as being a tobacco 
pathogen. The organism described by 
Delacroix might be the same as Pseudo- 
monas polycolor. Braun and Elrod 
(Jour. Bact., 4S, 1942, 40) are of the 
opinion that Clara's pathogen is Pseudo- 
monas aeruginosa. 

Description taken from Clara (Cornell 
Agr. Exp. Sta. Mem. 159, 1934, 28). 

Rods : 0.75 to 1 .2 by 1.05 to 3.0 microns. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Beef-extract agar colonies: Gra3ash- 
white, circular, raised, thin transparent 
margins. 

Broth: Turbid in 36 hours with thin 
pellicle. 

Milk: Alkaline; no curd. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 5^, 1942,601). 

Acid but not gas from glucose, galac- 
tose, fructose, mannose, arabinose, xy- 
lose, mannitol and glycerol. Alkaline 
reaction from salts of acetic, citric, malic, 



FAMILY PSEUDOMONADACEAE 



127 



lactic and formic acid. Rhamnose, su- 
crose, maltose, lactose, raffinose and sali- 
cin not fermented . 

Starch not hydrolyzed. 

Facultative anaerobe. 

Good growth in broth plus 7 per cent 
salt. 

Optimum temperature 25" to 30°C. 
Maximum 37° to 39°C. 

Distinctive character: Differs from 
Pseudomonas mellea in tj'pe of lesion pro- 
duced, does not digest starch, nor reduce 
nitrates and does not form acid from lac- 
tose nor sucrose. Pathogenic for labora- 
tory animals (Elrod and Braun, Sci. 94, 
1941, 520). 

Source : Repeatedly isolated from leaf 
spot of tobacco in the Philippines. 

Habitat: Pathogenic on tobacco. 

99. Pseudomonas viridiflava (Burk- 
holder) Clara. {Phytomonas viridiflava 
Burkh., Cornell Agr. Exp. Sta. Mem. 127, 
1930, 63; Clara, Science, 75, 1934, 111; 
Bacterium viridiflavum Burgwitz, Phyto- 
pathogenic Bacteria, Leningrad, 1935, 
127.) From Latin viridis, green; flaviis, 
yellow. 

Description from Clara (Cornell Agr. 
Exp. Sta. Mem. 139, 1934, 30). 

Rods: 0.75 to 1.5 by 1.5 to 3.15 microns. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Beef -extract agar colonies : Grayish- 
white, margins corrugated, edges irregu- 
lar. 

Broth: Turbid in 36 hours. 

Milk: Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H2S produced. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 82, 1942, 601). 

Acid but not gas from glucose, fruc- 
tose, mannose, arabinose, xylose, manni- 
tol and glycerol. Alkaline reaction from 
salts of acetic, citric, malic, lactic and 



succinic acids. Sucrose, lactose, mal- 
tose, raffinose, salicin, and salts of formic 
and tartaric acids not fermented. 

Starch: No hydrolysis. 

Growth in broth plus 5 per cent NaCl. 

Facultative anaerobe. 

Source: Two cultures isolated from 
spotted beans, one from England and 
one from Switzerland. 

Habitat : Pathogenic on bean, Phaseo- 

lus vidgaris. 

99a. Pseudomonas viridiflava var. con- 
centrica (Petersen) corrib. nov. {Phyto- 
monas viridiflava var. concentrica Peter- 
sen, Tridsskr. f. Planteavl., 88, 1932, 851; 
Bacterium viridiflavum var. conceniricum 
Burgwitz, Phytopathogenic Bacteria, 
Leningrad, 1935, 127.) From M. L. 
concentricus, concentric, referring to 
the rings on the colonies. 

Distinctive characters : Differs from 
Pseudomonas viridiflava in that it does 
not grow in Uschinsky's solution, and also 
in the shape of the colonies. 

Source : Isolated from the stems and 
leaves of blighted beans in Denmark. 

Habitat : Pathogenic on the bean, 
Phaseolus vidgaris. 

100. Pseudomonas ananas Serrano. 
(Serrano, Philipp. Jour. Sci., 55, 1934, 
355; Phytomonas ananas and Bacterium 
ananas Serrano, ibid, (not to be con- 
fused with Erwinia ananas Serrano, 
ibid., 86, 1928, 271); Bacterium serranoi 
Burgwitz, Bact. Dis. of Plants, Lenin- 
grad, 1936.) From Brazilian Indian, 
ananas, pineapple; M.L. Ananas, generic 
name . 

Rods : 0.6 by 1.8 microns. Motile with 
1 to 4 polar flagella. Gram -negative. 

Green fluorescent pigment produced 
in certain media. 

Gelatin: Liquefied. 

Beef-extract glucose agar colonies : 
White, v^ith undulating edges, smooth to 
rugose, glistening to dull. 

Beef -extract agar: Growth scant. 

Broth: Feeble growth. 



128 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Milk: Becomes alkaline with curd. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H2S formed. 

Acid but not gas from glucose, xylose 
and mannitol. Feeble with lactose. No 
acid with sucrose. 

Starch not hydrolyzed. 

Optimum temperature 30° to 31°C. 
Maximum 45°C. Minimum 7° to 10°C. 

Aerobic. 

Habitat : Causes a rot of pineapples, 
Ananas comosus. 

101. Pseudomonas ligustri (d'Oliveira) 
comb. nov. {Bacterium ligustri d'Oli- 
veira, Revista Agron., 24, 1936, 434.) 
From L. ligustrum, privet; M. L. Ligus- 
trum, a generic name. 

Rods: 0.5 to 0.7 by 1.3 to 3 microns. 
No chains. No capsules. Motile with 
2 to 5 polar flagella. Gram-negative. 

Green pigment produced on Dox agar, 
and in broth. 

Gelatin: Liquefied. 

Beef-extract agar colonies : Growth 
moderate. Milky white, circular, convex. 

Broth: Turbid ift 24 hours. No pel- 
licle. 

Milk : Coagulated in 6 days, and later 
digested. Litmus slightly acid. 

Nitrites not produced from nitrates. 

Indole not produced. 

Ammonia not produced. 

No gas from carbohydrates. Acid from 
glucose, galactose, arabinose and man- 
nose. No acid from sucrose, maltose, 
lactose, raffinose, mannitol and salicin. 

Source : From diseased Japanese privet 
in Lisbon, Portugal. 

Habitat: Pathogenic on privet, Ligtis- 
trum japonicum. 

102. Pseudomonas sesami Malkoff. 
(Malkoff, Cent. f. Bakt., II Abt., 16, 
1906, 665; Bacterium sesami Nakata, 
Ann. Phyt. Soc. Japan, 2, 1930, 242; 
Phytomonas sesami Kovachersky, Ann. 
Univ. de Sofia, Fac. Agron., 8, 1930, 464.) 
From Gr. sesamum, sesame; M. L. Sesa- 
mum, a generic name. 



Synonym: Nakata (loc. cit.) lists Bac- 
ierimn sesamicola Takimoto, Jour. Plant 
Protect. Tokyo, 8, 1927, 433 {Phyto- 
monas sesamicola Magrou, in Hauduroy 
et al.. Diet. d. Bact. Path., 1937, 412). 

Description from Nakata {loc. cit.). 

Rods: 0.6 to 0.8 by 1.2 to 3.8 microns. 
Motile with 2 to 5 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction rapid. 

Beef -agar colonies : Circular, flat, stri- 
ate, smooth, entire margins, white. 

Broth: Growth rapid. No pellicle. 

Milk: Alkaline. No coagulation. 

Nitrites not produced from nitrates. 

Indole not produced. 

No H2S produced. 

Acid but not gas from glucose. No 
acid from lactose, sucrose or glycerol. 

Starch not hydrolyzed. 

Optimum temperature 30°C. Maxi- 
mum 35°C. Minimum 0°C. 

Facultative anaerobe. 

Source : Isolated from brown spots on 
leaves and stems of sesame. 

Habitat: Pathogenic on sesame. 

103. Pseudomonas tolaasii Paine. 
(Paine, Ann. Appl. Biol., 6, 1919, 210; 
Phytomonas tolaasi Bergey et al., Man 
ual, 3rd ed., 1930, 259; Bacterium tolaasi 
Elliott, Bacterial Plant Pathogens, 1930, 
226.) Named for A. G. Tolaas who first 
reported the species. 

Rods: 0.4 to 0.5 by 0.9 to 1.7 microns. 
Motile with 1 to 5 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. , 

Gelatin: Liquefaction. 

Bouillon agar: Streak develops in 24 
hours, dirty bluish-white, wet-shining 
and slightly raised. 

Broth: Turbid in 24 hours. Pellicle. 

Milk: Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole production slight. 

Acid but not gas from glucose. No 
acid from lactose or sucrose. 



fa:mily pseudomoxadaceae 



129 



Starch hydroh-sis feeble. 

Optimum temperature 25°C. 

Source : Isolated in England from 
brown-spot of cultivated mushrooms. 

Habitat : Pathogenic on cultivated 
mushrooms. 

104. Pseudomonas xanthochlora 

(Schuster) Stapp. {Bacterium xantho- 
chlorum Schuster, Arbeit, a. d. Kaiserl. 
Biolog. Anstalt. f. Land. u. Forstw., 8, 
1912, 452; Phytomonas xanthochlora Ber- 
gey et al., Manual, 1st ed., 1923, 180; 
Stapp, in Sorauer, Handbuch der Pflan- 
zenkrankheiten, 2, 5 Auf., 1928, 213.) 
From Gr. xanthus, yellow; chlorus, green. 

Description from FJrw. Smith, Bacteria 
in Rel. to Plant Dis., 5, 1914, 272. 

Rods: 0.75 to 1.5 by 3.0 microns. Mo- 
tile with 1 to 3 fiagella. Gram-negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Slow liquefaction. 

Agar colonies : Circular, slightly raised, 
yellow -white. 

Broth: Strong clouding in 24 hours. 
A white pellicle. 

Milk : Slow coagulation and clearing. 

Nitrites are produced from nitrates. 

Indole is produced after 10 days. 

Hydrogen sulfide produced slowlj-. 

Acid but not gas from glucose and 
galactose. 

Optimum temperature 27"C. Maxi- 
mum 44°C. Minimum 2'C. 

Source: Isolated from rotting potato 
tubers in Germany. 

Habitat : Pathogenic on potato tubers 
and a number of unrelated plants. 

105. Pseudomonas rhizoctonia (Thom- 
as) comb. nov. {Aplanobacter rhizoctonia 
Thomas, Ohio Agr. Exp. Sta. Bull. 359, 
1922, 211 ; Bacterium rhizoctonia Stapp, 
in Sorauer, Handbuch der Pflanzenkrank- 
heiten, 2, 5 Auf., 1928, 290; Phytomonas 
rhizoctonia Burkholder, Ph3'topath., 20, 
1930, 7.) From Gr. rhizo, root; ctoiuis, 
murder. 

Rods: 0.5 to 0.85 by 1.4 to 1.9 microns. 
Non-motile. Gram-negative. 



Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Nutrient agar colonies: Greenish-yel- 
low, later olive-buff, circular, raised, 
slightly viscid. 

Broth: Turbid, pyrite yellow. 

Milk: Alkaline; clears. 

Nitrites are produced from nitrates. 

Indole reaction very slight. 

No HoS formed. 

Starch : Potato starch slighth- hy- 
drolyzed. 

Growth in 8 per cent salt. 

Optimum temperature 25° to 27''C. 
Maximum 38°C. Minimum 0°C. 

Source : Isolated from roots of lettuce 
showing the rosette disease. 

Habitat : Pathogenic on roots of let- 
tuce. 

106. Pseudomonas barker! (Berridge) 
Clara. (Bacillus of pear blossom dis- 
ease. Barker and Grove, Ann. Appl. Biol., 

I, 1914, 94; Barker and Grove's organism, 
Doidge, Ann. Appl. Biol., 4, 1917, 50; 
B. barkeri Berridge, Ann. Appl. Biol., 

II, 1924, 73; Phytomonas barkeri Bergey 
et al.. Manual, 3rd ed., 1930, 265; Bac- 
terium barkeri Elliott, Bacterial Plant 
Pathogens, 1930, 95; Clara, Science, 75, 
1934, 11.) Named for B. T. P. Barker 
who first reported the species. 

Description from Doidge (loc. cit.). 

Rods : 0.5 to 0.8 bj^ 2 to 4 microns. 
Motile with 1 to 4 polar flagella. Gram- 
negative (Burkholder), not Gram-posi- 
tive as stated. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Liquefaction. 

Agar: Growth is white, feeble, flat, 
glistening, smooth edged. 

Broth: Slightly turbid in 24 hours. 

Milk: Slowly cleared. 

Nitrites not produced from nitrates. 

Indole not formed unless culture 
warmed. 

Starch slowly digested. 

Source : Barker made many cultures 



130 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



from blighted pear blossoms. Doidge 
received a culture from Barker. 

Habitat: Causes a blossom blight of 
pear. 

107. Pseudomonas gladioli Severini. 
(Severini, Annali d. Bot., Rome, 11, 
1913, 420; Bacterium gladioli Elliott, 
Bact. Plant Pathogens, 1930, 132; Phyto- 
monas gladioli Magrou, in Hauduroy et 
al., Diet. d. Bact. Path., Paris, 1937, 
356.) From L. gladiolus, a little sword; 
M. L. Gladiolus, a generic name. 

Rods: 0.6 by 2.3 to 2.8 microns. Mo- 
tile with one or more polar flagella. 
Gram-negative. 

A pale yellow water-soluble pigment 
found, later orange. 

Gelatin colonies: Cream -colored, wart- 
like. Rapid liquefaction. 

Milk: Coagulated and slowly pep- 
tonized. 

Nitrites not produced from nitrates. 

Indole not formed. 

No gas. 

Aerobic. 

Optimum temperature 28° to 30°C. 

Habitat : Causes a corm rot of gladiolus 
and other tubers. 

108. Pseudomonas mellea Johnson. 
{Bacterium melleum Johnson, Jour. Agr. 
Res., ^S, 1923,489; Johnson, loc. cit., 489; 
Phytomonas mellea Bergey et al., Manual, 
3rd ed., 1930, 254.) From L. melleus, of 
or belonging to honey, the color of the 
colonies. 

Rods: 0.6 by 1.8 microns. Capsules. 
Motile with 1 to 7 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: Liquefied. 

Potato - glucose agar: Abundant 
growth, smooth, glistening, viscid, honey- 
colored. 

Broth: Turbid in 24 hours. Pellicle. 

Milk: Alkaline; clears. 

Nitrites not produced from nitrates. 

Indole not formed. 



No H2S formed. 

Starch hydrolysis feeble. 

Growth inhibited by 4 per cent salt. 

Optimum temperature, 26° to 28°C. 
Maximum 36°C. 

Facultative anaerobe. 

Distinctive character: Differs from 
Pseudomonas pseudozoogloeae in that it 
produces on tobacco a brown instead of a 
black spot with a halo, is orange-yellow 
in culture, and turns milk alkaline. 

Source: Isolated from brown rusty 
spots on tobacco in Wisconsin. 

Habitat : Pathogenic on leaves of to- 
bacco, Nicotiana tahacum. 

109. Pseudomonas betlis (Raguna- 
than) comb. nov. {Bacterium betle 
Ragunathan, Ann. Roy. Card., Pera- 
deniya, Ceylon, 11, 1928, 51; Aplanobac- 
ter betle Elliott, Bact. Plant Pathogens, 
1930, 4; Phytomonas betlis Magrou, in 
Hauduroy et al., Diet. d. Bact. Path., 
Paris, 1937, 337.) From Malayan, belle, 
betel, a kind of pepper, Piper betle. 

Rods: 0.5 by 1.5 to 2.5 microns, occur- 
ring singly or in short chains. Non- 
motile. Gram-negative. 

Green pigment formed in nutrient gela- 
tin and in broth. 

Gelatin: Liquefaction. 

Bovril agar colonies: Honey -yellow, 
circular at first, later echinulate. Raised, 
smooth and shiny. 

Broth: Surface becomes cloudy in 2 
days. Pellicle. 

No gas from lactose, maltose or sucrose. 

Starch is reduced. 

Aerobic. 

Source : Five cultures isolated from 
leaf spots on the betel vine. 

Habitat : Pathogenic on the betel vine, 
Piper betle. 

110. Pseudomonas panacis (Takimoto) 
Dowson. {Bacterium panaxi Nakata 
and Takimoto, Bui. Agr. Sta. Chosen, 5, 
1922, 1 ; Phytomonas panaxi Magrou, in 
Hauduroy et al.. Diet. d. Bact. Path., 
Paris, 1937, 389; Dowson, Trans. Brit. 



FAMILY PSEUDOMONADACEAE 



131 



Mycol. Soc, 26, 1943, 10.) From Gr. 
panax (panicis), a plant heal-all; M. L. 
Panax, a generic name. 

Description from Elliott, Bact. Plant 
Pathogens, 1930, 173. 

Rods : 0.5 by 1.3 to 1.5 microns. Chains. 
Motile with 4 to 6 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: Slight liquefaction. 

Agar colonies : White. 

Milk: Coagulated. 

No gas from sugars. 

Habitat : Causes a root rot of ginseng, 
Panax quinquefolium. 

111. Pseudomonas aleuritidis (McCul- 
loch and Demaree) Stapp. {Bacterium 
aleuritidis McCulloch and Demaree, 
Jour. Agr. Res., 45, 1932, 339; Stapp, Bot. 
Rev., 1, 1935, 408; Phytomonas aleuriti- 
dis Magrou, in Hauduroy et al.. Diet. d. 
Bact. Path., Paris, 1937, 328.) From 
Gr. aleurites, of wheaten flour; M. L. 
Aleurites, generic name. 

Rods: 0.6 to 0.7 by 1.1 to 3 microns. 
Motile with 1 to 5 polar, rarely bipolar, 
flagella. Capsules present. Gram-nega- 
tive. 

Green fluorescent pigment produced 
in certain media. 

Gelatin: Not liquefied. 

Beef agar slants: Growth is thin, white 
and viscid. 

Broth: A heavy white surface growth 
in 24 hours. Sediment. 

Milk: Becomes alkaline, but no sep- 
aration. 

Nitrites are produced from nitrates. 

Indole test feebly positive. 

Hydrogen sulfide test feebly positive. 

Acid but no gas from glucose, galac- 
tose and glycerol. Slow acid production 
from sucrose, maltose and lactose. 

Starch hydrolysis feeble. 

Optimum temperature 27" to 28°C. 
Maximum temperature 37°C. 

Optimum pH 6.2 to 6.8. pH range 5.4 
to 8.9. 



Source : Isolations from naturally in- 
fected tung oil trees in Georgia. 

Habitat: Pathogenic on the tung oil 
tree (Aleurites fordi), on the bean 
(Phaseolus vulgaris) and the castor bean 
(Ricinus communis). 

112. Pseudomonas giycinea Coerper. 
(Bacterium glycineum Coerper, Jour. 
Agric. Research, 18, 1919, 188; Coer- 
per, loc. cit., 188; Phytomonas giycinea 
Burkholder, Phytopath., 16, 1926, 922.) 
From glycys, sweet; ine, like; M.L. 
Glycine, generic name. 

Synonj-m : Bacterium sojae Wolf, Phy- 
topath., 10, 1920, 132 (Phytomonas sojae 
Burkholder, Phytopath., 16, 1926, 922; 
Pseudomonas sojae Stapp, in Sorauer, 
Handb. d. Pflanzenkrankheiten, 2, 5 
Aufl., 1928, 174). See Elliott, Bact. 
Plant Pathogens, 1930, 134; and Shunk 
and Wolf, Phytopath., 11, 1921, 18. 

Rods: 1.2 to 1.5 by 2.3 to 3 microns. 
Motile with polar flagella. Gram-nega- 
tive. 

Green fluorescent pigment produced in 
culture. 

Gelatin: Not liquefied. 

Beef-peptone agar colonies : Appear 
in 24 hours. Circular, creamy white, 
smooth, shining and convex. Margins 
entire. Butyrous in consistency. 

Milk: Litmus turns blue and later a 
separation of the milk occurs. Casein not 
digested. 

Nitrites not produced from nitrates. 

Indole test feebly positive. 

Not lipolytic (Starr and Burkholder, 
Phytopath., S2, 1942, 601). 

Starch not hydrolyzed. 

Acid from glucose and sucrose. 

Optimum temperature 24° to 26°C. 
Maximum 35°C. Minimum 2°C. 

Facultative anaerobe. 

Source : A number of cultures isolated 
from soy beans in Wisconsin. 

Habitat: Pathogenic on soybean. Gly- 
cine max (Soja max) . 

112a. Pseudomonas giycinea var. 
japonica (Takimoto) comb. nov. (Bac- 



132 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



terium soyae var. japonicum Takimoto, 
Jour. Plant Protect. Tokyo, U, 1927, 
556; Bacterium glycineum var. japonicum 
Elliott, Bact. Plant Pathogens, 1930, 136; 
Phytomonas glycinea var. japonica Ma- 
grou, in Hauduroy et al.. Diet. d. Bact. 
Path., Paris, 1937, 358.) From M. L. 
Japonicus, Japanese. 

Distinctive characters : Differs slightly 
from Pseudomonas glycinea in size of cell, 
length of chains, action in milk, and color 
in media. Okabe (Jour. Soc. Trop. 
Agr., Formosa, 5, 1933, 162) gives a de- 
scription of the organism which leads one 
to believe the differences are not great 
enough to be varietal. 

Source : Isolated from leaf spots on soy 
bean in Formosa. 

Habitat : Pathogenic on soy bean, Gly- 
cine max. 

113. Pseudomonas savastanoi (Erw 
Smith) Stevens. (Bacterium savastano 
Erw. Smith, U. S. Dept. Agr. Plant Ind 
Bull. 131, 1908, 31; Stevens, The Fungi 
which Cause Plant Diseases, 1913, 33 
Phytomonas savastanoi Bergey et al.. 
Manual, 1st ed., 1923, 190.) Named for 
F. Savastano, the Italian plant path- 
ologist. 

Note: Smith (loc. cit.) lists and dis- 
cards the following species since they were 
either mixed cultures or names with no 
descriptions : Bacterium oleae Arcangeli , 
Istit. Bot. delle R. Univ. di Pisa, Ri- 
cerche e Lavori, fasc. 1, 1886, 109; Bacil- 
lus oleae tuberculosis Savastano, Atti. 
R. Accad. Naz. Lincei Rend. CI. Sci. Fis., 
Mat. e Nat., 5, 1889, 92; Bacillus pril- 
lieuxianus Trevisan, I generi e le specie 
delle Batteriacee, Milano, 1889, 19; 
Bacillus oleae De Toni and Trevisan, in 
Saccardo, Sylloge Fungorum, 8, 1889, 
982. 

Description from Brown, Jour. Agr. 
Res., 44, 1932, 711. 

Rods: 0.4 to 0.8 by 1.2 to 3.3 microns. 
Motile with 1 to 4 polar flagella. Gram- 
negative. 

Green fluorescent pigment found in 
culture. 



Gelatin: No liquefaction. 

Beef agar colonies: White, smooth, 
flat, glistening, margins erose or entire. 

Broth : Turbid on the second day. No 
pellicle or ring. 

Milk: Becomes alkaline. 

Nitrites not produced from nitrates. 

No H2S produced. 

Acid but not gas from glucose, galac- 
tose and sucrose. 

Starch is hydrolyzed. 

Optimum temperature 23° to 24°C. 
Maximum 32°C. Minimum TC. 

Optimum pH 6.8 to 7.0. Maximum 8.5. 
Minimum 5.6. 

Aerobic. 

Source : Smith isolated his cultures 
from olive galls collected in California. 

Habitat: Pathogenic on olive. 

113a. Pseudomonas savastanoi var. 
fraxini (Brown) Dowson. (Bacterium 
savastanoi var. fraxini Brown, Jour. 
Agr. Res., 44, 1932, 721; Phytomonas 
savastanoi var. fraxini Magrou, in Hau- 
duroy et al., Diet. d. Bact. Path., Paris, 
1937, 410; Pseudomonas fraxini Skoric, 
Ann. Exp. For. Zagreb, 6, 1938, 66; Dow- 
son, Trans. Brit. Mycol. Soc, 26, 1943, 
11.) From M. L. Fraxinus, a generic 
name. 

Distinctive characters: Differs but 
slightly from Pseudomonas savastanoi, 
but is pathogenic on ash and not on olive. 

Source : Three cultures isolated from 
cankers on ash. 

Habitat: Pathogenic on ash, Fraxinus 
excelsior and F. americana. 

114. Pseudomonas tonelliana (Fer- 
raris) comb. nov. (Bacterium toncllianum 
Ferraris, Trattato di Patologia e Terapia 
Vegetale, 3rd ed., 1, 1926, 104; Phytomo- 
nas tonelliana Adams and Pugsley, Jour. 
Dept. Agr. Victoria, 32, 1934, 304.) 
Named for Tonelli, the Italian botanist. 

Synonym : Pseudomonas savastanoi 
var. ncrii C. O. Smith, Phytopath., 18, 
1928, 503. 

Description from Smith (loc. cit.) un- 
less otherwise noted 



FAMILY PSEUDOMOXADACEAE 



133 



Rods: 0.5 to 0.6 by 1.5 to 2.5 microns. 
Motile with 1 to 3 polar flagella. Gram- 
negative (Adams and Pugsley, loc. cil.). 

Gelatin: No liquefaction. 

Potato glucose agar colonies: Flat, 
circular, shining, margins somewhat un- 
dulated. 

Broth: Dense clouding with partial 
pellicle. 

Milk: Alkaline. No separation. 

Nitrites not produced from nitrates 
(Adams and Pugsley). 

Indole produced. Not produced (Adams 
and Pugsley). 

Acid but not gas from glucose and 
sucrose. No acid from lactose (Adams 
and Pugsley). 

Starch not hydrolyzed (Adams and 
Pugsley). 

Distinctive character : Pseudomonas 
savastanoi is similar in culture but is not 
pathogenic on oleanders. 

Source: Both Ferraris and C. O. Smith 
isolated the pathogen from galls on olean- 
der. 

Habitat: Pathogenic on oleander, 
Nerium oleander. 

115. Pseudomonas calendulae (Taki- 
moto) Dowson. {Bacleriuin calendulae 
Takimoto, Ann. Phytopath. Soc. Japan, 
5, 1936, 341; Phytomonas calendulae 
Burkholder, in Manual, 5th ed., 1939, 
201; Dowson, Trans. Brit. Mycol. Soc, 
26, 1943, 9.) From Latin, calendae, 
throughout the month; M.L. Calendula, 
a generic name. 

Rods: 0.5 by 1 to 2 microns. Motile 
with 1 to 3 polar flagella. Gram-nega- 
tive. 

Green fluorescent pigment produced 
in Uschinsky's and in Cohn's solutions. 

Gelatin: Not liquefied. 

Agar colonies: Circular, smooth, fiat, 
dirty white. 

Broth: Turbid. 

Milk: No coagulation. 

Nitrites not produced from nitrates. 

Indole formed in small amount. 

No HiS produced. 



Acid but not gas from glucose and 
glycerol. No acid from lactose or sucrose. 

Starch not hydrolyzed. 

Optimum temperature 27° to 30°C. 
Maximum 37°C. Minimum 0° to 7°C. 

Habitat : Pathogenic on marigolds. 
Calendula officinalis. 

116. Pseudomonas cichorii (Swingle) 
Stapp. (Phytomonas cichorii Swingle, 
Phytopath., 15, 1925, 730; Stapp, in Sor- 
auer, Handbuchder Pflanzenkrankheiten, 
2, 5 Auf., 1928, 291; Bacterium cichorii 
Elliott, Bact. Plant Pathogens, 1930, 
112.) From Gr. cichoria, chicory; M. 
L. Cichoriiim, generic name. 

Probable synonyms : Pseudomonas en- 
diviae Kotte, Phyt. Ztschr., 1, 1930, 609; 
Phytomonas endiviae (Kotte) Clara, 
Cornell Agr. Exp. Sta. Mem. 159, 1934, 
26; and Bacterium formosaniim Okabe, 
Jour. Soc. Trop. Agr., Formosa, 7, 1935, 
65. 

Description from Clara (loc. cit.) which 
is a description of a culture of Pseudo- 
monas endiviae from Kotte. Swingle's 
description is verj^ meager. 

Rods : 0.75 to 1.5 by 1.5 to 3.75 microns. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: No liquefaction. 

Beef -extract agar colonies : Circular, 
grayish-white with bluish tinge, raised 
with slightly irregular edges. 

Broth : Turbid in 36 hours with a 
smooth viscous pellicle. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H2S formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but not gas from glucose, galac- 
tose, fructose, mannose, arabinose, xy- 
lose, mannitol and glycerol. Alkaline 
production from salts of acetic, citric, 
lactic, malic, succinic and tartaric acids. 



134 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Rhamnose, maltose, sucrose, lactose, raf- 
finose and salicin not utilized. 

Starch not hydrolyzed. 

Slight growth in broth plus 6 per cent 
NaCl. 

Optimum pH 6.8 to 7.1. Maximum 
9.2 to 9.4. Minimum 5.0 to 5.3 (Kotte, 
Phyt. Ztsch., 2, 1930, 453). 

Facultative anaerobe. 

Distinctive characters: Differs from 
Pseudomonas inlyhi in that it does not 
liquefy gelatin or reduce nitrates to ni- 
trites. 

Source : Isolated from rot of French 
endive, Cichorium intybus by Swingle 
and by Okabe, and from C. endivia by 
Kotte. 

Habitat : Pathogenic on endive, lettuce 
and larkspur. 

117. Pseudomonas cissicola (Taki- 
nioto) comb. nov. {Aplanobacter cissicola 
Takimoto, Ann. Phytopath. Soc. Japan., 
9, 1939, 43.) From Greek, cissus, ivy; 
M. L. Cissus a generic name. 

Rods: 0.5 to 0.9 by 1.0 to 2.0 microns. 
Non-motile. Capsules. Gram-negative. 

Green fluorescent pigment formed in 
Uschinsky's solution. 

Gelatin: No liquefaction. 

Potato-extract agar colonies: Circular, 
convex, smooth, and dirty white. 

Broth: Feeble clouding followed by 
precipitation of pellicle and rim. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not produced. 

No acid nor gas from sucrose, glucose, 
lactose and glycerol. 

Starch is not digested. 

Salt toleration is 3 per cent. 

Optimum temperature 30°C. Maxi- 
mum 35°C. Minimum lO'C. Thermal 
death point 49° to 50°C. 

Source : Isolated from black spots on 
leaves of Japanese ivy, Cissus japonica in 
Japan. 

Habitat : Pathogenic only on Cisstis 
japonica. 

118. Pseudomonas nectarophila 
(Doidge) Rosen and Bleeker. {Bac- 



terium nectarophila Doidge, Ann. Appl. 
Biol., 4, 1917, 73; Phytomonas nectaro- 
phila Bergey et al., Manual, 3rd ed., 
1930, 262 ; Rosen and Bleeker, Jour. Agr. 
Res., It6, 1933, 98.) From Gr. nectar, 
nectar; philus, loving. 

Rods: 0.5 to 0.7 by 0.6 to 1.5 microns. 
Motile with 1 to 5 polar flagella. Cap- 
sules. Gram-negative. 

Green fluorescent pigment produced 
in culture. 

Gelatin: No liquefaction. 

Nutrient agar colonies : Yellowish- 
white, wet-shining, smooth, margins ir- 
regular. 

Broth: Heavy turbidity in 24 hours. 
Sediment. 

Milk: Cleared. 

Nitrites not produced from nitrates. 

Indole not formed. 

Acid from glucose and galactose. No 
acid from sucrose. 

Starch hydrolysis feeble. 

Optimum temperature 25 to 30°C. 

Facultative anaerobe. 

Distinctive character: Differs from 
Pseudomonas barkeri in that it does not 
liquefy gelatin, nor produce indole. Pro- 
duces capsules. 

Source : Isolated from blighted pear 
blossoms in South Africa. 

Habitat: Pathogenic on pear blossoms. 

119. Pseudomonas viburni (Thorn- 
berry and Anderson) Stapp. {Phytomonas 
viburni Thornberry and Anderson, Phy- 
topath., 21, 1931, 912; Stapp, Bot. Rev., 
1, 1935, 407; Bacterium viburni Burgwitz, 
Phytopathogenic Bacteria, Leningrad, 
1935, 160.) From L. viburnum, the way- 
faring tree; M. L. Viburnum, a generic 
name. 

Rods: 0.5 to 1.0 by 1 to 2.0 microns. 
Capsules present. Motile with 2 to 4 
polar flagella. Gram-negative (Burk- 
holder) ; not Gram-positive as stated. 

Green fluorescent pigment produced in 
culture (Burkholder). 

Gelatin: No liquefaction. 

Glucose beef -extract colonies: Dull 
gray, circular, edges entire. 



FAMILY PSEUDOMONADACEAE 



135 



Broth: Turbid with pellicle. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H2S formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid from glucose and galactose, but 
not sucrose (Burkholder). 

Starch : No hydrolysis. 

Slight growth in 3.5 per cent salt 
(Burkholder). 

Optimum temperature 25°C. Mini- 
mum 12°C. Maximum 35°C. 

Aerobic. 

Source : Isolated from angular leaf 
spots and stem lesions on arrow-wood, 
Viburnujn opulus, etc. 

Habitat : Pathogenic on Viburnum spp. 

120. Pseudomonas mori (Bo3^er and 
Lambert) Stevens. {Bacterium vinri 
Boyer and Lambert, Compt. rend. Acad. 
Sci. Paris, 117, 1893, 342; Bacterium mori 
Boyer and Lambert emend. Erw. Smith, 
Science, 31, 1910, 792; Stevens, The Fungi 
which Cause Plant Diseases, 1913, 30; 
Bacillus mori Holland, Jour. Bact., 5, 
1920, 222; Phytomonas mori Bergey et 
al., Manual, 1st ed., 1923, 191.) From 
Gr. moriun, mulberry; ]\L L. Moms, a 
generic name. 

Synonyms: Elliott (Bact. Plant Path- 
ogens, 1930, 166) lists Bacillus cuboni- 
anus Macchiati, Staz. Sperim. Agr. Ital., 
S3, 1892, 228 (Macchiati described the 
disease due to Pseudomonas mori, but 
gave an incorrect description of the patho- 
gen) ; also Bacterium cubonianum Fer- 
raris, Curiano le Plante, 6, 1928, 180 
(Ferraris uses Macchiati 's name but the 
description of Pseudomonas ynori). 

Description from Smith {loc. cit.). 

Rods: 0.9 to 1.3 by 1.8 to 4.5 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Green fluorescent pigment produced in 
culture. 

Gelatin : Not liquefied. 

Agar colonies : White, slow -growing, 
smooth, flat, edges entire becoming un- 
dulate. 



Milk: Becomes alkaline and clears. 

Nitrites not produced from nitrates. 

Indole none or feeble production. 

Hydrogen sulfide not produced (Okabe, 
Jour. Soc. Trop. Agr., 5, 1933, 166). 

No growth in broth plus 4 per cent salt 
(Okabe, loc. cit.). 

No gas from carbohydrates. 

Temperature range 1°C to 35°C. 

Source: Smith isolated the pathogen 
from blighted shoots of mulberry from 
Georgia. Also received cultures from 
Arkansas and the Pacific Coast. 

Habitat : Pathogenic on mulberry, 
Morus. 

121. Pseudomonas stizolobii (Wolf) 
comb. nov. {Aplanobacter stizolobii Wolf, 
Phytopath., 10, 1920, 79; Bacterium stiz- 
olobii McCulloch, Phytopath., 18, 1928, 
460; Phytomonas stizolobii Bergey et al.. 
Manual, 3rd ed., 1930, 280.) From Gr. 
siizo, to prick; lobium, a little lobe; 
Stizolobium, a generic name. 

Rods: 0.6 to 0.7 by 1.0 to 1.6 microns. 
Non-motile (Wolf). Motile with a short 
polar flagellum (McCulloch). Capsules. 
Gram-negative. 

Gelatin : No liquefaction. 

Agar colonies : Circular, smooth, white, 
raised and opaque. Margins entire to 
slightly undulate. 

Broth: Slightly turbid throughout. No 
pellicle or ring. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

No acid or gas in peptone broth plus 
sugars. 

Starch not hydrolyzed. 

Optimum temperature 25° to 28°C. 

Distinctive characters : Differs from 
Pseudomonas sojae (Pseudomonas gly- 
cinea) in the smaller size of cell, and 
absence of pellicle and dense clouding of 
broth. The pathogen does not infect 
soy bean. 

Source : Isolated from the leaf spot of 
velvet bean. 

Habitat : Pathogenic on velvet bean, 
Stizolobium deeringianium. 



136 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



122. Pseudomonas viciae Uyeda. 
(Uyeda in Takimoto, Jour. Plant Pro- 
tect., Japan, 2, 1915, 845; Bacterium vi- 
ciae Nakata, see Elliott, Bact. Plant 
Pathogens, 1930, 259; Phytomonas viciae 
Magrou, in Hauduroy et al.. Diet. d. Bact. 
Path., Paris, 1937, 430.) From L. vicia, 
vetch; M. L. Vicia, a generic name. 

Rods: 0.5 to 0.8 by 1.2 to 2.0 microns. 
Motile with 2 to 4 polar flagella. Gram- 
positive. 

Green fluorescent pigment produced in 
culture. 

Gelatin colonies : Pale white, glisten- 
ing, finally turning brown. No liquefac- 
tion. 

Milk : Coagulates and clears. 

Nitrites not produced from nitrates. 

No H2S produced. 

Facultative anaerobe. 

Habitat : Pathogenic on the broad 
bean {Vicia faba), the turnip (Brassica 
rapa), the carrot (Daucus carota) and the 
sweet potato {Ipomoea batatas). 

123. Pseudomonas alliicola Burk- 

holder. (Burkholder, Phytopath., 32, 
1942, 146 ; Phytomonas alliicola Burk- 
holder, ibid.) From L. allium, onion; 
-cola, dweller. 

Rods: 0.7 to 1.4 by 1.05 to 2.8 microns. 
Motile with 1 to several polar flagella, at 
times bi -polar. Gram-negative. 

Gelatin : Liquefaction. 

Beef -extract peptone agar streaks : 
Moderate in growth, white at first, later 
dirty in appearance, edges wavy, consist- 
ency viscid. Medium deep brown. 

Potato-glucose agar frequently be- 
comes greenish. 

Broth: Turbid with light pellicle. 
Brown. 

Milk : Cleared and litmus reduced . 
Neutral. 

Nitrites produced from nitrates. 

Indole not produced. 

Hydrogen sulfide not produced. 

Lipolytic action very strong. 

Acid but no gas from 1-arabinose, d- 
xjdose, rhamnose, glucose, d-galactose, 
fructose, d-lactose, maltose, sucrose, 



glycerol, mannitol and salicin. Alkali 
from salts of acetic, citric, formic, hip- 
puric, lactic, malic, succinic, tartaric 
acids. 

Starch not hydrolysed. 

Slight growth in broth plus 4 per cent 
salt. 

Aerobic. 

Optimum temperature 30°C. Maxi- 
mum 41°C. Minimum 5°C. 

Source: Seven isolates from storage 
rot of onion bulbs. 

Habitat: Pathogenic on onion bulbs, 
Allium cepa. 

124. Pseudomonas gardeniae Burk- 
holder and Pirone. (Burkholder and 
Pirone, Phytopath., 31, 1941, 194; Phyto- 
monas gardeniae Burkholder and Pirone, 
ibid.) From M. L. Gardenia, a generic 
name. 

Rods: 0.75 by 2.4 microns. Motile 
with 1 to 2 polar flagella. Gram-nega- 
tive. 

Gelatin: Liquefaction. 

Beef -extract peptone agar colonies: 
Growth fair, white to dirty gray and vis- 
cid. Medium becoming dark brown. 

Potato-glucose agar : No brown color. 

Broth: Turbid with pellicle. Dark 
brown. 

Milk: Soft curd with pellicle. Clears 
in zones. Litmus reduced. 

Nitrites produced from nitrates. 

Hydrogen sulfide not produced. 

Indole not formed. 

Acid from glucose, galactose, xylose, 
rhamnose, sucrose, maltose, mannitol, 
glycerol, and salicin. Alkali produced 
from the salts of citric, malic, malonic, 
succinic, tartaric and hyppuric acids. 
Good growth in tyrosine and in aspara- 
gine broth. 

Starch is not hydrolyzed. 

Aerobic. 

Source: Eight isolates from leaf spots 
of gardenias in New Jersey. 

Habitat: Pathogenic on leaves of Gar- 
denia jasminoides. 

125 Pseudomonas caryophylli Burk- 
holder. (Burkholder, Phytopath., 31, 



FAMILY PSEUDOMOXADACEAE 



137 



1941, 143; Phytomonas caryophylli, 
Burkholder, ibid.) From M. L. Caryo- 
phyllus, an old generic name. 

Rods: 0.35 to 0.95 by 1.05 to 3.18 mi- 
crons. At times slightly curved. Motile 
with 1 to several polar flagella. Fre- 
quently bipolar. Gram-negative. 

Gelatin: Liquefaction after 3 to 4 
weeks. 

Potato glucose agar colonies: 3 to 4 
mm in diameter, circular, smooth, glis- 
tening, edges entire. Color is tan to gray 
mauve. Old culture dark brown. Con- 
sistency butj^rous. 

Broth: Turbid with a white sediment. 

Milk: Litmus slowly becomes blue. 
Slight reduction at bottom of tube. No 
clearing. 

Nitrites produced from nitrates. Also 
ammonia and gas are produced in a 
synthetic nitrate medium. Asparagine, 
KXO3 and XH4H2PO4 can be utilized. 

Indole not formed. 

Hydrogen sulfide not formed. 

Lipolytic action slight to moderate. 

Acid from 1-arabino.se, d-xylose, rham- 
nose, glucose, d-galactose, fructose, d- 
lactose, maltose, and sucrose, glj'cerol, 
mannitol, and salicin. Alkali with so- 
dium salts of acetic, citric, formic, hip- 
puric, lactic, malic, maleic, succinic and 
tartaric acid. 

Starch not hydrolyzed. 

Aerobic. 

Optimum temperature 30° to 33°C. 
Maximum 46°C. Minimum 5^C. or less. 

Slight growth in broth plus 3.5 per cent 
salt. 

Source: Isolated first by L. K. Jones 
and later by W. H. Burkholder from 
dying carnation plants from Spokane, 
Washington. Twelve isolates used in 
description. 

Habitat : Pathogenic on roots and 
stalks of the carnation, Dianthus caryo- 
phylliis. 

126. Pseudomonas solanacearum Erw. 

Smith. (Bacillus solanacearum Erw. 
Smith, U. S. Dept. Agr., Div. Veg. Phys. 
and Path., Bui. 12, 1896, 10; Bacterium 



solanacearum Chester, Ann. Kept. Del. 
Col. Agr. Exp. Sta., 9, 1897, 73; Erw. 
Smith, Bacteria in Relation to Plant 
Diseases, S, 1914, 178; Phytomonas so- 
lanacearum Bergej' et al., ^Manual, 1st 
ed., 1923, 186.) From L. solanum night- 
shade; AI. L. Solanaceae, a plant famih'. 

Probable synonyms: Elliott (Bact. 
Plant Pathogens, 1930, 203) lists the 
following: Bacillus nicotianae Uyeda, 
Cent. f. Bakt., II Abt., 13, 1904, 327; 
Bacillus sesami Malkoff and Pseudo- 
monas sesami Malkoff, Cent. f. Bakt., 
II Abt., 16, 1906, 664; Bacillus rnusae 
Rorer, Phytopath., 1, 1911, 45; Bacillus 
tnusaru?n Zeman, Rev. Facul. Agr. L'niv., 
La Plat, 14. 1921, 17; Erwinia nicotianae 
Bergey et al.. Manual, 1st ed., 1923, 172; 
Phytomonas ricini Archibald, Trop. Agr., 
Trinidad, 4, 1927, 124. 

Description taken from Elliott {loc. 
cit.). 

Rods: 0.5 to 1.5 microns. Motile with 
a polar flagellum. Gram-negative. 

Gelatin: Xakata (Jour. Sci. Agr. Soc. 
Tokyo, 294, 1927, 216) states there are 
two forms, one of which shows slight 
liquefaction. The other shows no lique- 
faction. 

Agar colonies: Small, irregular, round- 
ish, smooth, wet-shining, opalescent, 
becoming brown. 

Broth: Slight pellicle. Broth turns 
brown . 

]Milk : Cleared without precipitation 
of casein. 

Nitrites produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not produced (Burk- 
holder). 

Glucose, sucrose, glycerol, sodium 
citrate, peptone, tyrosine, asparagine and 
glutanic acid are utilized (Mushin, Aus- 
tral. Jour. Expt. Biol, and Med., 16, 
1938, 325). 

Nitrogen sources utilized are ammonia, 
nitrates (KNO3) asparagine, tyrosine, 
peptone and glutamic acid, but not potas- 
sium nitrite (Mushin, loc. cit.). 

Starch not hydrolyzed. 



138 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Optimum temperature 35° to 37°C. 
Maximum 41 °C. Minimum 10°C. 

Pathogenicity readily lost in culture. 

Source : Isolated from brown-rot of 
solanaceous plants. 

Habitat : Soil pathogen in warm moist 
climates attacking numerous species of 
plants, especially potato, tobacco, and 
tomato. . 

126a. Pseudomonas solanacearum 
var. asiatica (Erw. Smith) Stapp. {Bac- 
terium solanacearum var. asiaticum Erw. 
Smith, Bact. in Relation to Plant Dis- 
eases, 3, 1914, 282; Stapp, in Sorauer, 
Handbuch der Pflanzenkrankheiten, 2, 
5 Auf., 1928, 253; Phytomonas solana- 
cearum var. asiatica Magrou, in Hau- 
duroy et al., Diet. d. Bact. Path., Paris, 
1937, 414.) From Gr. asiaticus, asiatic. 

Distinctive characters: Differs from 
Pseudomonas solanacearum in that it 
turns litmus milk and cream red. 

Source : Isolated by J. A. Honing from 
diseased tobacco plants in Medan, Suma- 
tra. 

127. Pseudomonas castaneae (Kawa- 
mura) comb. nov. {Bacterium castaneae 
Kawamura, Ann. Phytopath. Soc. Japan, 
S, 1934, 15; Phytomonas castaneae Ma- 
grou, in Hauduroy et al.. Diet. d. Bact. 
Path., Paris, 1937, 343.) From M. L. 
Castanea, a generic name. 

Rods: 0.8 to 1.2 by 1.0 to 1.8 microns. 
Motile with 1 to 5 polar flagel la. Gram- 
negative . 

Gelatin: Liquefied. 

Beef agar colonies : White, circular, 
edges slightly undulate, viscid. 

Milk: No coagulation. Peptonized. 

Acid but not gas from glucose, sucrose 
and glycerol. No acid from lactose. 

Optimum temperature 25° to 27°C. 
Maximum 35°C. Minimum 3°C. 

Facultative anaerobe. 

Habitat : Causes water-soaked spotting 
on leaves and shoots of chestnut, Cas- 
tanea. 

128. Pseudomonas seminum Cayley. 
(Cayley, Jour. Agr. Sci., 8, 1917, 461; 
Bacterium seminum Stevenson, Foreign 



Plant Dis., U.S.D.A. Office of Sec'y-, 
1926, 141 ; Phytomonas seminum Bergey 
et al., Manual, 3rd ed., 1930, 272.) From 
L. semen {seminus) seed. 

Rods : 1 .0 by 4.0 to 5.0 microns. Spore- 
like bodies present. Capsules. Motile 
with a single flagellum. Gram-positive. 

Gelatin: Rapid liquefaction. 

Agar colonies: White, more or less 
circular, transparent, spreading. 

Broth: Turbid. Pellicle. 

Litmus milk: Milk becomes clear and 
apricot color. 

Nitrites produced from nitrates. 

Acid but not gas from glucose and 
sucrose. No acid from lactose. 

Starch : No hydrolysis. 

Optimum temperature 25°C. 

Facultative anaerobe. 

Source : Isolated from seeds, stems and 
pods of diseased peas in England. 

Habitat : Pathogenic on peas. 

129. Pseudomonas passiflorae (Reid) 
comb. nov. {Phytomonas passiflorae Reid, 
New Zealand Jour. Sci. and Tech., 22, 
1939, 264a.) From L. passio, passion; 
flora, flower; M. L. Passiflora, a generic 
name. 

Rods: 0.2 to 0.5 by 1.2 to 3.2 microns. 
Motile with 1 to 5 polar flagella. Cap- 
sules present. Gram-negative. 

Gelatin : Liquefied. 

Beef-peptone agar colonies : Small, flat, 
smooth, dry, shining, translucent, gray- 
ish and butyrous. 

Broth : Turbid in 4 days. Transient 
pellicle. 

Milk: Slightly alkaline. No coagula- 
tion nor clearing. 

Nitrites not produced from nitrates. 
No growth on synthetic nitrate agar. 

Indole not formed. 

Hydrogen sulfide not formed. 

Acid reaction occurs in galactose, starch 
and sucrose. No gas. 

Starch is not hydrolysed. 

Source : From diseased leaves and fruit 
of the passion-fruit in New Zealand. 

Habitat : Pathogenic on Passiflora 
edulis. 



FAMILY PSEUDOMONADACEAE 



139 



130. Pseudomonas fabae (Yu) comb, 
nov. {Phytomonas fabac Yu, Bull, of the 
Chinese Bot. Soc, 2, 1936, 34.) From 
Latin, faba bean. 

Rods: 0.8 to 1.1 by 1.1 to 2.8 microns. 
Motile with 1 to 4 polar flagella. Gram- 
negative. 

Gelatin: Liquefied. 

Nutrient agar colonies : Circular, en- 
tire, viscid, glistening, raised, smooth to 
wrinkled, white to salmon. Medium 
amber. 

Broth: Turbid after 12 hours. Pellicle. 

Milk : Growth slow. Clears. 

Nitrites produced from nitrates. 

Indole production slight. 

Hydrogen sulfide not produced. 

Acid but not gas from glucose. No 
acid nor gas developed from arabinose, 
xylose, fructose, galactose, sucrose, 
lactose, maltose, raffinose, dextrin, in- 
ulin, mannitol or adonitol in a 1 per cent 
Bacto-peptone broth. 

Starch: Verj^ weak diastatic action. 

Optimum temperature 35°C. ^laxi- 
mum 37 to 38"C. Minimum 4°C. Ther- 
mal death point 52 to 53°C. 

Aerobic. 

Growth retarded in 2 per cent salt. 
Very slight growth in 3 per cent salt. 

Source: From diseased broad beans 
at Nanking, China- 
Habitat : Pathogenic on broad or Wind- 
sor bean, Vicia faba. 

131. Pseudomonas astragali (Taki- 
moto) comb. nov. {Bacterium astragali 
Takimoto, Jour. Plant Protect., 17, 1930, 
732; Phytomonas astragali Burkholder, 
in Manual, 5th ed., 1939, 197.) From 
M. L. Astragalus, a generic name. 

Description translated by Dr. K. 
Togashi. 

Rods: 0.7 to 0.8 by 1.2 to 2.2 microns. 
Motile, with 1 or 2 flagella. Gram- 
negative. 

Gelatin: Liquefied. 

Agar plates: Growth somewhat slow, 
colorless or grayish-white, entire margins, 
more or less aqueous, butyrous. 



Uschinsky's medium : Growth vigorous, 
turbid, not viscid, ring, and sediment. 

Milk: No coagulation of casein, slow 
digestion. Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced in small 
amount. 

No acid or gas from glucose, sucrose, 
lactose and glycerol in broth. 

Starch not hydrolyzed. 

Temperature relations : Minimum be- 
low 5° and maximum 33'^C. Thermal 
death point 50° to 51 °C. 

Aerobic. 

Source : Species isolated from Astra- 
galus sp. 

Habitat : Causes a black leaf -spot of 
Astragahis sp. 

132. Pseudomonas colurnae (Thorn- 
berry and Anderson) comb. nov. {Phyto- 
monas colurnae Thornberry and Ander- 
son, Phytopath., 27, 1937, 948.) From 
the species, Corylus colurna. 

Rods: 0.8 to 1.0 by 1.0 to 1.8 microns. 
Single, in pairs or chains. Capsules. 
Motile with 1 to 2 polar flagella. Gram- 
negative. 

Gelatin: Liquified. 

Glucose agar slants: Growth filiform, 
raised, dull, smooth, opaque and viscid. 

Broth : Moderate turbidity. Ring. 

Milk : Peptonization complete with 
acid production. No reduction of litmus 
nor coagulation. 

Nitrites not produced from nitrates. 

Indole not produced. 

Hydrogen sulfide not produced. 

No appreciable amount of gas from 
xylose, glucose, sucrose or glycerol. 

Starch hydrolyzed. 

Optimum temperature 21°C. Mini- 
mum 5°C. Maximum 35°C. Thermal 
death point oO^C. 

Aerobic. 

Source : From leaves and young stems 
of the Turkish hazelnut in Illinois. 

Habitat : Pathogenic on the Turkish 
hazelnut, Coryltis colurna. 



140 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



133. Pseudomonas maublancii (Foex 
and Lansade) comb. nov. (Bacterium 
maublancii Foex and Lansade, Comp. 
rend. Acad. Sci. Paris, 202, 1936, 2174; 
Phytomonas maublancii Burkholder, in 
Manual, 5th ed., 1939, 198.) Named for 
M. Maublanc, French colonial plant 
pathologist. 

Rods : 0.4 by 1.3 microns. Motile with 
1 to 3 polar flagella. Gram-negative. 

Gelatin: Liquefied. 

Gelatin colonies: Round, translucent, 
margins entire. 

Broth: Thin pellicle. 

Milk: Not coagulated; clears. 

Nitrites not produced from nitrates. 

Indole not formed. 

No HoS formed. 

Carbohydrates not fermented. 

Ammonia produced. 

Growth in Fermi's solution, not in Us- 
chinsky's solution. 

Source : Isolated from rotting vascular 
and parenchymatic tissue of banana 
stalks. 

Habitat: Causes a disease of the 
banana plant. 

134. Pseudomonas polygon! (Thorn- 
berry and Anderson) coynb. nov. (Phyto- 
monas polygoni Thornberry and Ander- 
son, Phytopath., 27, 1937, 947.) From 
Gr. polygonum, knot-weed; M. L. Poly- 
gonum, a generic name. 

Rods: 0.5 to 1.5 by 1.5 to 2.5 microns. 
Motile with 2 to 8 bi -polar flagella. Cap- 
sules. Gram-positive (?). Other spe- 
cies reported by these investigators as 
Gram-positive have proved to be Gram- 
negative on a retest (Burkholder). 

Gelatin: Liquified. Brown. 

Glucose agar slant: Abundant, fili- 
form, flat, dull, smooth, pale olive-gray, 
butyrous. Medium turns brown. 

Broth: Turbid. Pellicle. 

Milk: Alkaline and clears. Litmus 
not reduced. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not produced. 



No appreciable amount of gas from car- 
bohydrates. 

Starch : No hydrolysis. 

Optimum temperatures 18°C. Mini- 
mum 7°C. Maximum 35°C. 

Aerobic. 

Source : From diseased leaves of Poly- 
gonum convolvulus in Illinois. 

Habitat : Pathogenic on black bind- 
weed, Polygonum convolvulus. 

135. Pseudomonas iridicola (Taki- 
moto) Stapp. (Bacterium iridicola Taki- 
moto. Fungi, Nippon Fungological Soc, 
1, 1931, 24; Stapp, Bot. Rev., 1, 1935, 408; 
Phytomonas iridicola Burkholder, in 
Manual, 5th ed., 1939, 198.) From Gr. 
iris (iridis) , iris, a rainbow; -cola, 
dweller. 

Rods: 0.7 to 0.8 by 1.2 to 2 microns. 
Motile with 1 to 3 polar flagella. Gram- 
negative. 

Gelatin: Liquefied. 

Beef agar colonies: White, circular, 
raised or convex. 

Milk: Clears without coagulation. 

No acid or gas from carbohydrates. 

Starch digested. 

Optimum temperature 38°C. Mini- 
mum 4°C. 

Source : Isolated from a brown leaf 
spot of iris. 

Habitat : Pathogenic on Iris tectorum 
and Iris japonica. 

136. Pseudomonas levistici Oster- 
walder. (Ostcrwalder, Cent. f. Bakt., 
II Abt., 25, 1909, 260; Bacterium levistici 
Stevenson, Foreign Plant Dis., U. S. 
Dept. Agr., Oflfice of Sec'y-, 1926, 101; 
Phytomonas levistici Magrou, in Hau- 
duroy et al.. Diet. d. Bact. Path., Paris, 
1937, 373.) From M. L. Levisticum, a 
generic name. 

Rods: 0.5 to 0.7 by 1.1 to 1.5 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: Colonies greenish-white. 
Liquefaction. 

Nutrient agar: Good growth at room 
temperature. Yellowish-white. 



FAMILY PSEUDOMONADACEAE 



141 



Broth: Pellicle. 

Indole formed. 

No H-iS produced. 

Source: Isolated from spots on the 
leaves of lovage. 

Habitat: Pathogenic on lovage, Levis- 
ticum officinale. 

137. Pseudomonas radiciperda (Jav- 
oronkova) Stapp. {Bacterium radici- 
perda Javoronkova, Bull. Plant Protect., 
Leningrad, Ser. II, .5, no. 1, 1932, 161; 
Stapp, Bot. Rev., 1, 1935, 408; Phyto- 
monas radiciperda Magrou, in Hauduroy 
et al., Diet. d. Bact. Path., 1937, 401.) 
From L. radix (radicis), root; pcrdo, to 
destroy. 

Description from Javoi'onkova, Rev. 
App. Myc, 11, 1932, 652. 

Rods : O.S by 1 to 2 microns. Capsules. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Gelatin: Liquefaction. 

Beef-peptone agar colonies: Round, 
smooth, shining, white to pale yellow. 

Milk: Peptonized. 

Indole not formed. 

No H-jS formed. 

Acid but not gas from carbohydrates. 

Optimum temperature 23° to 25°C. 

Aerobic. 

Habitat : Causes a root rot of red clover 
(Trifolium pratense), lentils (Lens escu- 
lenta) and lucerne. 

138. Pseudomonas melophthora Allen 
and Riker. (Allen and Riker, Phyto- 
path., 22, 1932, 557; Bacterium meloph- 
thorum Allen and Riker, ibid.; Phyfo- 
monas melophthora Allen and Riker, 
ibid.) From Gr. melum, apple; phthora, 
destroyer. 

Rods: 0.68 by 1.32 microns. Motile 
with 2 polar flagella. Gram-negative; 
Gram-positive cells appear in old cultures. 

Gelatin : No liquefaction. 

Nutrient agar plus 2 per cent glucose: 
Colonies appear in 36 hours. After 3 
days colonies circular, smooth, glistening, 
convex; edges entire; light pink, but not 
constant. 



Broth : Good growth. Pellicle and 
sediment. 

Milk : Little change, if an.y. 

Nitrites not produced from nitrates. 

Indole not formed. 

No HoS produced. 

Acid from arabinose, glucose, galac- 
tose, fructose, sucrose and glycerol. No 
acid from lactose, maltose, dextrin and 
inulin. 

Starch not hydrolyzed. 

Optimum temperature 21° to 25°C. 

Source : Description based on 7 cul- 
tures isolated from rotting apples and 
from apple maggots. 

Habitat : Pathogenic on apples, and 
found with the apple maggot, Rhagoletis 
pomonella. 

139. Pseudomonas helianthi (Kawa- 
mura) comb. nov. (Bacterium helianthi 
Kawamura, Ann. Phyt. Soc. Japan, 4, 
1934, 27; Phytomonas helianthi Magrou, 
in Hauduroy et al., Diet. d. Bact. Path., 
Paris, 1937, 362.) From M. L. Helian- 
thus, a generic name. 

Probable synonym : Phytomonas helian- 
thi var. tubero.si Thornberry and Ander- 
son, Phytopath., 27, 1937, 948. 

Rods: 1 to 1.4 by 1.6 to 2.4 microns. 
Motile with a single polar flagellum. 
Gram-negative. 

Gelatin: No liquefaction. 

Beef agar colonies: White, circular, 
edges entire. 

Broth: Turbid. Pellicle. 

Milk: Peptonized. Litmus reduced. 

Nitrates: Gas production. 

Indole not produced. 

No HoS produced. 

Acid but not gas from sucrose and glyc- 
erol. No acid from lactose and maltose. 

Starch hydrolyzed. 

Optimum temperature 27° to 28°C. 
Maximum 35.5°C. Minimum 12°C. 

Good growth at pH 6.4. No growth 
pH 5.4 and pH 8.8. 

Habitat : Pathogenic on sunflower, 
Helianthus debilis. 

140. Pseudomonas alboprecipitans 
Rosen. (Rosen, Ann. Missouri Bot. 



142 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Garden, 9, 1922, 383; Bacterium albo- 
precipitans Elliott, Bact. Plant Path., 
1930, 89; Phytomonas alboprecipitans 
Bergey et al., Manual, 3rd ed., 1930, 
277.) From Latin albus, white and 
precipitans , precipitating, referring to 
the white precipitate produced in cul- 
ture. 

Rods: 0.6 by 1.8 microns, occurring 
singly or in pairs. Capsules present. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: Not liquefied. 

Nutrient agar colonies : White, circular, 
raised, smooth, sticky, with margins 
entire. Whitish discoloration of the 
medium. 

Broth: Turbid in 24 hours. Heavy 
sediment in old cultures. 

Milk : Becomes alkaline and slowly 
clears. 

Nitrites produced from nitrates. 

Indole not produced. 

No HzS produced. 

Acid but not gas from glucose, fruc- 
tose, glycerol and mannitol. No acid 
from lactose, maltose or sucrose. 

Starch is hydrolyzed. 

Optimum temperature 30° to 35°C. 
Maximum temperature 40 °C. Minimum 
0°C. 

Aerobic. 

Distinctive characteristics : White pre- 
cipitate in culture media. 

Source : Isolated a number of times 
from foxtail grass. 

Habitat : Pathogenic on foxtail, Chaeto- 
chloa lutescens and other grasses. 

141. Pseudomonas petasitis (Taki- 
moto) comb. nov. {Bacterium petasitis 
Takimoto, Ann. Phyt. Soc. Japan, 2, 
1927, 55; Phytomonas petasitis Magrou, 
in Hauduroy et al., Diet. d. Bact. Path., 
Paris, 1937, 393.) From M. L. Petasites, 
a generic name. 

Rods: 0.8 to 1.1 by 1.1 to 1.7 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: No liquefaction. 



Beef agar colonies : White, circular or 
ameboid, butyrous. 

Broth: Strong turbidity. Pellicle. 

Milk: Coagulated in 30 days. 

Nitrites produced from nitrates with 
gas formation. 

Indole not produced. 

No HoS produced. 

No evident acid in peptone broth but 
gas from glucose, lactose and sucrose. 
.\cid but not gas from glycerol. 

Weak growth in broth plus 6 percent 
salt. 

Optimum temperature 27° to 30 °C. 
Maximum 47°C. Minimum approxi- 
mately 5°C. 

Source: Isolated from brown to black 
lesions on Petasites japonicus in Japan. 

Habitat : Pathogenic on leaves of 
Petasites japonicus. 

142. Pseudomonas lignicola Wester- 
dijk and Buisman. (De lepenziekte, 
Arnliem, 1929, 51.) From Latin, lignum, 
wood; -cola, dweller. 

Rods: Single or short chains. Motile 
with 1 to several polar flagella. Gram- 
negative. 

Gelatin: No liquefaction. 

Malt agar streaks : Milk white with a 
colorless edge. 

Broth : Turbid with light pellicle. 

Milk: No coagulation. No acid. 

Nitrites not produced from nitrates. 

Indole not formed. 

Starch hydrolysis slight. 

Optimum temperature ±25°C. 

Source : From vessels of elm wood 
showing dark discoloration, in Holland. 

Habitat: Pathogenic in elm wood. 

143. Pseudomonas andropogoni (Erw. 
Smith) Stapp. {Bacterium andropo- 
goni Erw. Smith, Bacteria in Relation 
to Plant Diseases, 2, 1911, 63; Elliott and 
Smith, Jour. Agr. Res., 38, 1929, 4; Stapp, 
in Sorauer, Handbuch der Pflanzen- 
krankheiten, 2, 5 Auf., 1928, 27; Phyto- 
monas andropogoni Bergey et al., 
Manual, 3rd ed., 1930, 276.) From M. 



FAMILY PSEUDOMONADACEAE 



143 



L. Andropogon , a generic name (a syn- 
onym of Holcvs). 

Description from Elliott and Smith 
(loc. cit.). 

Rods: 0.64 by 1.76 microns. Motile 
with one to several bipolar flagella. 
Capsules. Gram-negative. 

Gelatin: Feeble liquefaction or none. 

Beef -extract agar colonies : Slow -grow- 
ing, round, smooth, glistening, viscid, 
white. 

Broth : Growth slow with moderate 
turbidity in 48 hours. A thin pellicle. 

Milk: Alkaline and clears. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H2S formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but not gas from glucose, arabi- 
nose, fructose and xylose. No acid from 
.sucrose, maltose, lactose, raffinose, glyc- 
erol and mannitol. 

Starch partially digested. 

Optimum temperature 22' to 30°C. 
Ma.ximum 37° to 38 °C. Minimum 1.5°C. 

Optimum pH 6.0 to 6.6. Maximum 
8.3 to 8.6. Minimum 5.0. 

Source : Elliott used for her description 
4 cultures isolated from lesions on sorgo, 
sorghum and broom-corn. 

Habitat : Pathogenic on sorghum, Hol- 
cus sorghum. 

144. Pseudomonas woodsii (Smith) 
Stevens. {Bacterium woodsii Erw. 
Smith, Bact. in Relation to Plant Dis- 
eases, 2, 1911, 62; Stevens, Plant Disease 
Fungi, New York, 1925, 39; Phytomonas 
woodsii Bergey et al., Manual, 3rd ed., 
1930, 256.) Named for A. F. Woods, 
American plant pathologist. 

Description from Burkholder and Gu- 
terman, Phytopath., 25, 1935, 118. 

Rods: 0.67 by 1.56 microns. Motile 
with a polar flagellum. Gram-negative. 

Gelatin: No liquefaction. 

Beef -extract agar slants : Growth slow 
and scant, filiform, creamy, butyrous. 

Broth: Turbid. 



Milk: Becomes alkaline but otherwise 
little changed. 

Nitrites not produced from nitrates. 

Indole not formed. 

No HoS formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 601). 

Acid but not gas from glucose, fruc- 
tose, galactose, arabinose, xylose, rham- 
nose, lactose, glycerol and mannitol. 
Alkaline reaction from salts of acetic, 
citric, malic and succinic acids. Sucrose, 
maltose, salicin, and lactic and formic 
acids not fermented. Starch not hy- 
drolyzed. 

Slight growth in broth plus 3 per cent 
salt. 

Aerobic. 

Source : Isolated from water-soaked 
lesions on carnation leaves. 

Habitat: Pathogenic on carnation, 
Dianthus caryophyllvs. 

145. Pseudomonas panici-miliacei 
(Ikata and Yamauchi) comb. nov. {Bac- 
terium panici-jriiliacei Ikata and Yamau- 
chi, Jour. Plant Protect., 18, 1931, 35; 
Phytomonas panici-miliacei Burkholder, 
in Manual, 5th ed., 1939, 204.) From 
M. L. Panicum miliaceum. 

Description translated by Dr. K. 
Togas hi. 

Rods: 0.8 to 1.1 by 1.8 to 2.6 microns. 
Motile, with a single flagellum. Gram- 
negative. 

Gelatin : Not liquefied. 

Potato-agar plates : Growth moderate, 
whitish, then tinged with light orange, 
undulating margins. 

Broth: Turbid, white pellicle formed. 

Milk: No coagulation and slow diges- 
tion. Alkaline. 

Nitrites are produced from nitrates. 

Indole not formed. 

No H2S produced. 

No acid and no gas from sucrose, glu- 
cose, lactose, glycerol and sodium 
nitrate. 

Starch not hydrolyzed. 

Optimum temperature 30° to SS'C. 



144 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Facultative anaerobe. 

Source : Species first isolated from 
millet, Panicum miliaceum. 

Habitat : Causes a leaf stripe of Pani- 
cum miliaceum. 

146. Pseudomonas saliciperda Lindei- 
jer. (Lindeijer, Inaug. Diss., Univ. 
Amsterdam, 1932; Phytopath. Ztschr., 
6, 1933, 373; Bacterium saliciperda 
Burgwitz, Phytopathogenic Bacteria, 
Leningrad, 1935, 106; Phytomonas salici- 
perda Magrou, in Hauduroy et al., Diet, 
d. Bact. Path., Paris, 1937, 408.) From L. 
salix (salicis), willow; perdo, to destroy. 

Rods: 1.2 to 2.1 microns in length. 
Motile with a polar fiagellum. Gram- 
negative. 

Gelatin: No liquefaction. 

Beef wort agar colonies: Gray -white. 

Milk: No acid nor coagulation. 

Nitrites produced (small amount) 
from nitrates. 

Indole formation slight. 

No gas from carbohydrates. 

Starch not hydrolyzed. 

Facultative anaerobe. 

Source: Isolated from wilted branches 
of willow and pathogenicity proved. 

Habitat : Pathogenic on willow, Salix 
spp. 

147. Pseudomonas eriobotryae (Taki- 
moto) Dowson. (Bacterium eriobotryae 
Takimoto, Jour. Plant Protect., 18, 1931, 
354; Phytomonas eriobotryae Burkholder, 
in Manual, 5th ed., 1939, 205; Dowson, 
Trans. Brit. Mycol. Soc, 26, 1943, 10.) 
From M. L. Eriobotrya, a generic name. 

Translated by Dr. K. Togashi. 

Rods: 0.7 to 0.9 by 2.2 to 3.0 microns. 
Motile, w^ith 1 or 2 flagella. Gram- 
negative. 

Gelatin: Not liquefied. 

Agar-plates : Colonies appear after 3 
days, white or hyaline, butyrous, margins 
entire. 

Broth: Moderately turbid, pellicle 
powdery, ring formed. 



Milk : No coagulation, peptonized 
slowly. Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

No H^S produced. 

No acid or gas from glucose, sucrose, 
lactose and glycerol in broth. 

Starch not hydrolyzed. 

Temperature relations : Minimum 
below 4°C, optimum 25° to 26°C, and 
maximum 32°C. Thermal death point 
51°C. 

Aerobic. 

Source : Species isolated from loquat, 
Eriobotrya japonica. 

Habitat : Causes a bud rot of Eriobotrya 
japonica. 

148. Pseudomonas wieringae (Elliott) 
comb. nov. {Phytomonas betac Wieringa, 
Nederl. Tijdschr. Hyg., Microbiol, en 
Serol., Leiden, 2, 1927, 148; Bacterium 
wieringae Elliott, Man. Bact. Plant 
Pathogens, 1930, 264; Phytomonas wier- 
ingae Burkholder, in Manual, 5th ed., 
1939, 206.) Named for K. L. Wieringa, 
plant pathologist of Holland. 

Because Bacterium betae Chester (Ann. 
Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 
53) may be a pseudomonad, the species 
name proposed by Elliott has been 
retained. 

Description from Elliott (loc. cit.). 

Rods : 0.5 to 2.0 microns. Motile with 
1 to 5 polar flagella. Gram-negative. 

Beef-agar colonies: Smooth, round, 
white to grayish, fluorescent. 

Milk: Cleared in 5 days. Not co- 
agulated. 

Nitrites not produced from nitrates. 

No gas from sugars. 

Optimum temperature 28° to 30 °C. 
Maximum 37°C. Minimum 4°C. 

Source : Isolated from vascular rot of 
beets in Holland. 

Habitat : Pathogenic on beets, Beta 
vulgaris. 



FAMILY PSEUDOMONADACEAE 



145 



Appendix I*: The following species are 
believed to belong in the genus Pseudo- 
monas although descriptions are fre- 
(luently incomplete. 

Achromohacter pellucidinn Harrison. 
(Canadian Jour. Res., 1, 1929, 236.) 
Isolated from halibut. For a description 
of this species, see Bergey et al., Manual, 
othed., 1939, 619. 

Bacillus aurantiacus tingitanus Rem- 
linger and Bailly. (jNIaroc Medical, 
No. 150, 1935; See Lasseur, Dupaix and 
Babou, Trav. Lab. Microbiol. Fac. 
Pharm. Nancy, Fasc. 8, 1935, 41.) From 
water. Dissociates readily. Related to 
Pseudomonas fluorescens aureus Zim- 
mermann. See p. 645. 

Bacillus cyaneofluorescens Zaugemeis- 
ter. (Cent, f . Bakt., I Abt., 18, 1895, 321 ; 
Pseudomonas cyaneofluorescens Migula, 
Syst. d. Bakt., 2, 1900, 906.) From blue 
milk. 

Bacillus fluorescens nivalis Eisenberg. 
(Eine Gletscherbakterie, Schmelck, 
Cent. f. Bakt., 4, 1888, 545; Eisenberg, 
Bakt. Diag., 3 Aufl., 1891, 77.) From 
the melting snow of a glacier. Probably 
a S3'nonym of Pseudomonas fluorescens. 
Bacillus lactis saponacei Weigmanii 
and Zirn. (Cent. f. Bakt., 15, 1894, 468. * 
From soapy milk. 

Bacterium auxinophilum Jacobs. 
(Ann. Appl. Biol., 23, 1935, 619.) A 
Gram-negative organism with a polar 
flagellum which liquefies gelatin rapidly. 
Bacterium bosporum Kalnins. (Lat- 
vijas Universitates Raksti, Serija I, 
No. 11, 1930, 259.) Decomposes cellu- 
lose. Single polar flagellum. From soil. 
Bacterium briosii FsLvavino. (Atti 1st. 
Bot. d. R. Univ. di Pavia, Ser. 2, 12, 
1910, .337.) The natural host is Lyco- 
persicon esculentum. Pavarino (Rev. di 
Patol. Veg., 6, 1913, 161) states that this 
organism and Phytobacter lycopersicuni 
Groenewege (Meded. Rijks. Hoogere 
Land, Tuin- en Boschbouwschool, Dell 5, 
5, 1912, 217) should be considered identi- 



cal. It closely resembles Bacterium 
vesicatoriuin Doidge (Jour. Dept. Agr. 
So. Africa, 1, 1920, 718) according to 
Gardner and Kendrick (Jour. Agr. Res., 
21, 1921, 140). 

Bacterium daphorum Kalnins. (Lat- 
vijas Universitates Raksti, Serija I, 
No. 11, 1930, 257.) Decomposes cellu- 
lose. Single polar flagellum. From soil. 
Bacterium fraenkelii Hashimoto. (Zeit. 
f. Hyg., 31, 1899, 88.) A pleomorphic 
polar flagellate bacterium. From milk. 
Bacterium gummis Comes. (Comes, 
Napoli, Maggio 18, 1884, 14; see Comes, 
Atti d. R. 1st. d'incoraggiamento alii 
Sc, Ser. 3, 3, 1884, 4; Bacillus gummis 
Trevisan, I generi e le specie delle 
Batteriacee, Milano, 1889, 17.) Patho- 
genic on grapes, Viiis spp. 

Bacterium krameriani Pavarino. 
(Atti R. Accad. Naz. Lincei Rend. CI. 
Sci. Fis., Mat. et Nat., 20, 1911, 233.) 
Pathogenic on the orchid, Oncidium 
krameriani. 

Bacterium pusiolum Kalnins. (Lat- 
vijas U'niversitates Raksti, Serija I, 
No. 11, 1930, 261.) Decomposes cellu- 
lose. Single polar flagellum. From 
manure. 

Bacterium protozoides Kalnins. (Lat- 
\'ijas U'niversitates Raksti, Serija I, 
No. 11, 1930, 263.) Decomposes cellu- 
lose. Single polar flagellum. From soil. 
Pseudomonas acuta INIigula. (Culture 
No. 11, Lembke, Arch. f. Hyg., 29, 1897, 
317 ; Migula, Syst. d. Bakt., ~2, 1900, 921 .) 
From the intestine. 

Pseudomonas alba Migula. {Bacillus 
fluorescens albus Zimmermann, Bakt. 
unserer Trink- u. Nutzwasser, I Reihe, 
1890, 18; Migula, Syst. d. Bakt., 2, 1900, 
909.) From water. Bacillus fluorescens 
non liquefaciens Eisenberg, Bakt. Diag., 
3 Aufl., 1891, 145 may be identical ac- 
cording to Migula {loc. cit.). 

Pseudomonas allii (Griffiths) Migula. 
{Bacterium allium Griffiths, Proc. Roy. 
Soc. Edinburgh, 51, 1887, 40; Migula. 



* Appendixes I and II prepared by Prof. Robert S. Breed, New York State Experi- 
ment Station, Geneva, New York, July, 1943. 



146 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Syst. d. Bakt., 2, 1900, 932.) From 
rotted onions. 

Pseudomonas aquatilis Migula. (Tata- 
roff, Inaug. Diss., Dorpat, 1891, 31; 
Migula, Syst. d. Bakt., 2, 1900, 933.) 
From water. Said to form spores. 

Pseudomonas aromatica Migula. 
(Bacillus crassus aromaticus Tataroff, 
Inaug. Diss., Dorpat, 1891, 27; Migula, 
Syst. d. Bakt., 2, 1900, 880.) From 
water. 

Pseudomonas aromatica var. quercilo- 
pyrogallica Kluyver, Hof and Boe- 
zaardt. (Enzymologia, 7 1939, 28.) 

Pseudomonas arliculaia Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 
462.) From the stomachs of birds. 

Pseudomonas aucubicola Trapp. 
(Phytopath., 26, 1936, 264.) Isolated 
from Aucuba japonica. Not pathogenic. 

Pseudomonas aurea Migula. (Bacillus 
fluorescens aureus Zimmermann, Bakt. 
unserer Trink- u. Nutzwasser, I Reihe, 
1890, 14; Migula, Syst. d. Bakt., 2, 1900, 
931.) From water. 

Pseudomonas brassicae acidae G ruber. 
(Cent. f. Bakt., II Abt., 22, 1909, 558). 
From sauerkraut. Identical with Bac- 
terium brassicae acidae Conrad (Arch. f. 
Hyg., 29, 1897, 75) according to Lehmann 
and Neumann (Bakt. Diag., 5 Aufi., 2 
1912, 380.) 

Pseudomonas bulyri Migula. (Bacil- 
lus bulyri fluorescens Lafar, Arcli. f. 
Hyg., 13, 1891, 19; Migula, Syst. d. Bakt., 
2, 1900, 894.) From butter. 

Pseudomonas calciphila Molisch. 
(Cent. f. Bakt., II Abt., 65, 1925, 136.) 
From fresh water. Deposits CaCOa. 

Pseudomonas calco-acetica Clifton. 
(Enzymologia, 4, 1937, 246.) 

Pseudomonas capsulata Migula. 
(Syst. d. Bakt., 2, 1900, 915; not Pseudo- 
monas capsulata Bergey et al., Manual, 
1st ed., 1923, 124.) Similar to Pseudo- 
monas macroselmis Migula. 

Pseudomonas caryocyanea (Dupaix) 
Beijerinck. (Bacillus caryocyaneus Du- 
paix, Thesis, Univ. of Nancy, 1933, 1 ; 
Beijerinck, see Dupaix', ibid., 13; Bac- 



terium caryocyaneum Dupaix, ibid., 246.) 
Isolated from rotten willow wood, from 
yeast mash and beer-wort. Name ap- 
pears first as Bacillus caryocyaneus on a 
culture sent by Beijerinck from Delft, 
Holland to the National Collection of 
Type Cultures, Lister Institute, London. 
Regarded by Dupaix as closely related to 
Bacillus cyaneo -fluorescens Zangemeister 
(Cent. f. Bakt., I Abt., 18, 1895, 321; 
Pseudomonas cyaneo -fluorescens Migula, 
Syst. d. Bakt., 2, 1900, 906); Der blaue 
Bacillus, Mildenberg (Cent. f. Bakt., 
II Abt., 56, 1922, 309; Pseudomonas 
mildenbergii Bergey et al., Manual, 3rd 
ed., 1930, 172); and Bacillus pyocyaneus 
Gessard (Compt. rend. Acad. Sci. Paris, 
94, 1882,536). 

Pseudovionas catarrhalis Chester. 
(Bacillus der Hundestaupe, Jess, Cent, f . 
Bakt., II Abt., 25, 1899, 541; Chester, 
Man. Determ. Bact., 1901 , 308.) Isolated 
from catarrh in dogs. 

Pseudomonas caviae Scherago. (Jour. 
Bact., 31, 1936, 83; Jour. Inf. Dis., 60, 
1937, 245.) Cause of an epizootic septi- 
cemia in guinea pigs. 

Pseudomonas chlorophaena Migula. 
(Syst. d. Bakt.,^, 1900, 899.) 

Pseudomonas coccacea Migula. (Cul- 
ture No. 10, Lembke, Arch. f. Hyg., 29, 
1897, 317; Migula, Syst. d. Bakt., 2, 
1900,924.) From the intestine. 

Pseudomonas cohaerea (sic) (Wright) 
Chester. (Bacillus cohaerens Wright, 
Mem. Nat. Acad. Sci., 7, 1895, 464; 
Chester, Man. Determ. Bact., 1901, 
312.) From water. 

Pseudomonas coli Migula. (Culture 
No. 8, Lembke, Arch. f. Hyg., 29, 1897, 
315; Migula, Syst. d. Bakt., 2, 1900, 920.) 
From the intestine. 

Pseudomonas colloides Migula. 
(Bacillus fluorescens putidus colloides 
Tataroff, Inaug. Diss., Dorpat, 1891, 40; 
Migula, Syst. d. Bakt., 2, 1900, 902.) 
From water. Said to form spores. 

Pseudomonas conradi Lehmann and 
Neumann. (Bakt. Diag., 5 Aufl., 2, 
1912, 394.) Red pigment. 



FAMILY PSEUDOMONADACEAE 



u: 



Pseudonionas delabcns (Wright J 
Chester. (Bacillus delabens Wright, 
Mem. Nat. Acad. Sci., 7, 1895, 456; 
Chester, Man. Determ. Bact., 1901, 
314.) From water. 

Pseudonionas duplex Migula. (Cul- 
ture No. 7, Lembke, Arch. f. Hyg., 29, 
1897, 314; Migula, Syst. d. Bakt., 2, 
1900,922.) From the intestine. 

Pseudotnonas cllipsoiden Migula. 
(Bacillus oogenes fluorcsccns 0, Zorken- 
dorfer, Arch. f. Hyg., 16, 1893, 393; 
Migula, Syst. d. Bakt., 2, 1900, 925.) 
From hens' eggs. 

Pseudomonas ephemerocyanea Fuller 
and Norman. (Jour. Bact., Ji.6, 1943, 
274.) From soil. Decomposes cel- 
lulose. 

Pseudomonas erylhra Fuller and Nor- 
man. (Jour. Bact., 46, 1943, 276.) From 
soil. Decomposes cellulose. 

Pseudomonas erythrospora (Cohn) 
Migula. (Bacillus erylhrosporus Cohn, 
Beitr. z. Biol. d. Pfianzen, S, Heft 1, 
1879, 128; Migula, in Engler and Prantl, 
Die nautrl. Pflanzenfam., 1, la, 1895, 29.) 
From air, meat infusion and water. 
Said to form spores. 

Pseudomonas fimhriata (Wright) 
Chester. (Bacillus fimbriatus Wright, 
Mem. Nat. Acad. Sci., 7, 1895, 463; 
Chester, Man. Determ. Bact., 1901, 
313.) From water. 

Pseudomonas fluorcsccns cxiliosus van 
Hall. (Ztschr. f. Pflanzenkr., IS, 1903, 
132.) Causes soft rot of shoots and bulbs 
of iris (Iris spp.). 

Pseudomonas foliacea Chester. 
(Bacillus fluorcsccns foliaceus Wright, 
Mem. Nat. Acad. Sci., 7, 1895, 439; 
Chester, Man. Determ. Bact., 1901, 324; 
Bacillus fluorescens-foliaceus Chester, 
ibid.) From water. Very similar to 
Pseudomonas incognita Chester. 

Pseudomonas gasoformans ]\Iigula. 
(Ein neuer gasbildender Bacillus, Gart- 
ner, Cent. f. Bakt., 15, 1894, 1; Migula, 
Syst. d. Bakt., 2, 1900, 883.) Gas 
bubbles in gelatin stab. 



Pseudomonas gracilis Migula. (Syst. 
d. Bakt., 8, 1900, 888.) Morphologically 
like Pseudomonas fluorescens Migula. 

Pseudomonas granulata Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 
464.) From the stomach and intestine 
of birds. 

Pseudomonas halestorgus Elazari- 
Volcani. (Ph.D. Thesis, Hebrew Univ., 
Jerusalem, 1940.) A halophilic pseudo- 
monad from the Dead Sea. 

Pseiido7no7ias hydrosulfurea Migula. 
(Bacillus oogenes hydrosidfureus fi, Zor- 
kendorfer, Arch. f. Hyg., 16, 1893, 385; 
Migula, Syst. d. Bakt., 2, 1900, 898.) 
From hens' eggs. 

Pseudomonas iridis van Hall. (Van 
Hall, Thesis, Univ. Amsterdam, 1902 and 
Ztschr. f. Pflanzenkr., 13, 1903, 129; 
Bacterium iridis Elliott, Man. Bact. 
Plant Path., 1930, 142; Phytomonas iridis 
Magrou, in Hauduroy et al., Diet. d. 
Bact. Path., Paris, 1937, 369.) Causes a 
rot of bulbs and leaves of iris (7ns spp.). 

Pseudoinonas iris (Frick) Migula. 
(Bacillus iris Frick, Arch. f. path. Anat., 
116, 1889, 292; according to Eisenberg, 
Bakt. Diag., 3 Aufl., 1891, 148; Migula, 
Syst. d. Bakt., ^,1900,931.) 

Pseudomonas ilalica (Foa and Chia- 
pella) Reinelt. (Quoted from Lehmann 
and Neumann, Bakt. Diag., 7 Aufi., 2, 
1927, 367.) Phosphorescent. 

Pseudomonas javanica (Eijkmann) 
Migula. (Photobacterium javanense 

Eijkmann, Geneesk. Tijdschr. v. 
Nederl. -Indie, 32, 1892, 109; Abst. in 
Cent. f. Bakt., 12, 1892, 656; Bacillus 
java7iiensis Dyar, Annals New York 
Acad. Sci., 8, 1895, 359; Bacterium java- 
niensis Chester, Man. Determ. Bact., 
1901, 170; Photobacterium javanicum 
Lehmann and Neumann, Bakt. Diag., 1 
Aufl., 2, 1896, 199; Migula, Syst. d. Bakt., 
2, 1900, 953.) From sea fish in Java. 
Blue-green luminescence. 

Pseudomonas lactica Weiss. (Arb. bakt. 
Inst. Karlsruhe, 2, Heft 3, 1902, 238.) 
From a vegetable infusion. 

Pseudomonas lasia Fuller and Norman. 



148 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



fJour. Bact., 46, 1943, 275.) From soil. 
Decomposes cellulose. 

Pseudomonas lembkei Migula. (Cul- 
ture No. 12, Lembke, Arch. f. Hyg., 29, 
1897, 318; Migula, Syst. d. Bakt., 2, 1900, 
896.) From the intestine. 

Pseudomonas liquefaciens (Tataroff) 
Migula. {Bacillus liquefaciens Tataroff, 
Inaug. Diss., Dorpat, 1891, 29; Migula, 
Syst. d. Bakt., 2, 1900, 876.) From water. 

Pseudomonas listeri Weiss. (Arb. bakt. 
Inst. Karlsruhe, 2, Heft 3, 1902, 260.) 
From a vegetable infusion. 

Pseudomonas Zong^a Migula. {Bacillus 
fluorescens longus Zimmermann, Bakt. 
unserer Trink- u. Nutzwasser, I Reihe, 
1890, 20; Migula, Syst. d. Bakt., 2, 1900, 
907.) From water. 

Pseudomonas macruselmis Migula. 
{Bacillus Jlvorescens putidus Tataroff, 
Inaug. Diss., Dorpat, 1891, 42; Migula, 
in Engler and Prantl, Die natlirl. Pflan- 
zenfam., 1, la, 1895, 29.) From water. 

Pseudomonas maidis (Eisenberg) 
Migula. {Bacillus maidis Eisenberg, 
Bakt. Diag., 3 Aufl., 1891, 119; Migula, 
Syst. d. Bakt., 2, 1900, 877.) From corn 
grains soaked in water and from feces 
of pellagra patients. 

Pseudomonas maschekii Migula. 
(Blaugriiner Bacillus, Maschek, Bakt. 
Untersuch. d. Leitmeritzer Trinkwasser, 
Jahresber. d. Oberrealschule zu Leit- 
meritz, 1887; Migula, Syst. d. Bakt. 2, 
1900, 916.) From water. 

Pseudomonas melochlora (Winkler and 
Schrotter) Migula. {Bacillus melo- 
chloros Winkler and Schrotter, Ein neuer 
grunen Farbstoff entwickelnder Bacillus, 
Wien, 1890; Migula, Syst. d. Bakt., 2, 
1900, 893.) From caterpillar feces. 

Pseudomonas mesenterica Migula. 
{Bacillus fluorescens mesentericus Tata- 
roff, Inaug. Diss., Dorpat, 1891, 38; 
Migula, Syst. d. Bakt., 2, 1900, 903.) 
From water. 

Pseudomonas metalloides Migula. 
{Bacillus rosaceus metalloides Tataroff, 
Inaug. Diss., Dorpat, 1891, 65; not Bac- 
terium rosaceus metalloides Dowdeswell, 
Ann. de Microgr., 1, 1888-89, 310, see 



Heffernan, Cent. f. Bakt., II Abt., 8, 
1902, 689; Pseudomonas rosacea Migula, 
in Engler and Prantl, Die natlirl. Pflan- 
zenfam., 1, la, 1895, 29; Migula, Syst. d. 
Bakt., 2, 1900, 938.) Single flagellum. 
Red and yellow -red pigment. From 
water. 

Pseudomonas minutissima Migula. 
{Bacillus fluorescens liquefaciens 
minutissimus Unna and Tommasoli, 
Monatsh. f. prakt. Dermat., 8, 1889, 57; 
according to Eisenberg, Bakt. Diag., 3 
Aufl., 1891, 76; Migula, Syst. d. Bakt., 2, 
1900, 891.) Found on human skin in 
cases of seborrheic eczema. 

Pseudomonas mohilis Migula. (Cul- 
ture No. 9, Lembke, Arch. f. Hyg., 29, 
1897, 316; Migula, Syst. d. Bakt., 2, 
1900,923.) From the intestine. 

Pseudomonas monadiformis (Kruse) 
Chester. {Bacillus coli mohilis Messea, 
Riv. d'Igiene, Rome, 1890; Bacillus 
monadiformis Kruse, in Fltigge, Die 
Mikroorganismen, 2, 1896, 374; Chester, 
Man. Determ. Bact., 1901, 308.) From 
typhoid stools. 

Pseudomonas mucidolens Levine and 
Anderson. (Jour. Bact., 2S, 1932, 337.) 
Causes musty odors in eggs. Also milk 
(Olsen and Hammer, Iowa State Coll. 
Jour. Sci., 5, 1934,125). 

Pseudomonas mucidolens var. tarda 
Levine and Anderson. (Jour. Bact., 23, 
1932, 337.) Causes mustj'- odors in eggs. 

Pseudomonas nexibilis (Wright) 
Chester. {Bacillus nexibilis Wright, 
Mem. Nat. Acad. Sci., 7, 1895, 441; 
Chester, Man. Determ. Bact., 1901, 309.) 
From water. 

Pseudomonas nivalis Szilvinyi. 
(Cent. f. Bakt., II- Abt., 94, 1936, 216.) 
A red chromogen isolated from red snow 
in Austria. 

Pseudomonas ochroleuca Migula. 
{Bacillus y, Zorkendorfer, Arch. f. Hyg., 
16, 1893, 396; Migula, Syst. d. Bakt., 2, 
1900, 897.) From hens' eggs. 

Pseudomonas oogenes Migula. {Bacil- 
lus oogenes hydrosidfureus 8, Zorken- 
dorfer, Arch. f. Hyg., 16, 1893, 386; 



FAMILY PSEUDOMONADACEAE 



149 



Migula, Syst. d. Bakt., 2, 1900, 878.) 
Single flagellum. From hens' eggs. 

Pseudomonas ovi Migula. (Bacillia^ 
oogenes fluorescens e, Zorkendorfer, Arch, 
f. Hyg., 16, 1893, 395; Migula, Syst. d. 
Bakt., 2, 1900, 924.) From hens' eggs. 

Pseudomonas ovicola Migula. {Bacil- 
lus oogenes fluorescens y, Zorkendorfer, 
Arch. f. Hyg., 16, 1893, 394; Migula, 
Syst. d. Bact., 2, 1900, 925.) From hens' 



Pseudomonas pallescens Migula. 
(Bacillus viridis pallescens Frick, in 
Virchow, Arch. f. path. Anat., 116, 1889, 
292; according to Eisenberg, Bakt. Diag., 
3 Aufl., 1891, 154; Migula, Syst. d. Bakt., 
2, 1900, 927.) Source not given. 

Pseudomonas pansinii Migula. 
(Bacillus fluorescens non liquefaciens 
Pansini, in Vixchow, Arch. f. path. Anat., 
122, 1890, 452; Migula, Syst.-d. Bakt., 2, 
1900, 926.) 

Pseudomonas pelliculosa Migula. 
(Bacillus oogenes fluorescens d, Zorken- 
dorfer, Arch. f. Hyg., 16, 1893, 395; 
Migula, Syst. d. Bakt., 2, 1900, 926.) 
From hens' eggs. 

Pseudomonas pellucida Kern. (.Arb. 
bakt. Inst. Karlsruhe, /, Heft 4, 1896, 
465.) From the intestine of birds. 

Pseudomonas plehniae Spieckermann 
and Thienemann. (Arch. f. Hyg., 74, 
1911, 110.) Isolated from carp. Patho- 
genic for many species of fish. 

Pseudomonas plicala (Frankland and 
Frankland) Migula. (Bacillus plicatus 
Frankland and Frankland, Phil. Trans. 
Roy Soc. London, 178, B, 1887, 273; 
Migula, Syst. d. Bakt., 2, 1900, 881.) 
From air. 

Pseudomonas pseudolyphosa Migula. 
(Typhusahnlicher Bacillus, Lustig, 
Diag. d. Bakt. d. Wassers, 1893, 16; 
Migula, Syst. d. Bakt., 2, 1900, 893.) 
From water. 

Pseudomonas pidlulans (Wright) 
Chester. (Bacillus pullidans Wright, 
Mem. Nat. Acad. Sci., 7, 1894, 445; 
Chester, Man. Determ. Bact., 1901, 315.) 
From water. 



Pseudomonas prolea Frost. (U. S. 
PubHc Health Ser., Hyg. Lab. Bull. 66, 
1910, 27.) From filtered river water. 
Can be agglutinated by specific typhoid 
immune-serum. 

Pseudomonas rosea Chester. (Bacil- 
lus roseus vini Bordas, Joulin and 
Rackowski, Compt. rend. Acad. Sci. 
Paris, 126, 1898, 1550; Chester, Man. 
Determ. Bact., 1901, 327; not Pseudo- 
monas rosea Migula, in Engler and Prantl, 
Dienaturl.Pfllanzenfam.,i, la, 1895,30.) 
From wine. Said to form spores. 

Pseudomonas (Hydrogenomonas) sac- 
charophila Doudoroff. (Enzymologia, 
9, 1940, 50.) From stagnant water. 

Pseudomonas sapolactica (Eichholz) 
De Rossi. (Bacterium sapolacticum 
Eichholz, Cent. f. Bakt., II Abt., 9, 
1902, 631; De Rossi, Microbiologia 
Agraria e Technica, Torino, 1927, 693.) 
Isolated from soapy milk. 

Pseudomonas sencea Migula. (Seiden- 
glanzender Bacillus, Tataroff, Inaug. 
Diss., Dorpat, 1891, 26; Migulu, Syst. d. 
Bakt., 2, 1900, 882.) 

Pseudomonas tenuis Migula. (Bacil- 
lus fluorescens tenuis Zimmermann, Bakt. 
unserer Trink- u. Nutzwasser, I Reihe, 
1890, 16; Migula, Syst. d. Bakt., 2. 
1900,910.) From water. 

Pseudomonas Irommehchldgel (Rav- 
enel) Chester. (Bacillus iromnielschldgel 
Ravenel, INIem. Nat. Acad. Sci., 8, 1896, 
26; Chester, Man. Determ. Bact., 1901, 
327.) From soil. Said to form spores. 

Pseudomonas undidala Chester. 
(Bacillus fluorescens unditlalus Ravenel, 
Mem. Nat. Acad. Sci., 8, 1896, 20; 
Chester, Man. Determ. Bact., 1901, 
328.) From soil. Said to form spores. 

Pseudomonas virescens (Frick) Migula. 
(Bacillus virescens Frick, Arch. f. path. 
Anat., 116, 1889, 292; Migula, Syst. d. 
Bakt., ^,1900,916.) From green sputum. 

Pseudomonas viridans (Symmers) 
Migula. (Bacillus viridans Symmers, 
Brit. Med. Jour., No. 1615, 1891, 1252; 
Abst. in Cent. f. Bakt., 12, 1892, 165; 
Migula, Syst. d. Bakt., 2, 1900, 890.) 
From water. 



150 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Psctidorunnas viridescens Chester. 
(Bacillus viridescens liquefaciens Rav- 
enel, Mem. Nat. Acad. Sci., 8, 1896, 24; 
Chester, Man. Determ. Bact., 1901, 328.) 
From soil. Said to form spores. 

Pseudomonas viridis Migula. (Bacil- 
lis der grimen Diarrhoe der Kinder, 
Lesage, Arch. d. Physiol, norm, et path., 
20, 1888, 212; see Eisenberg, Bakt. Diag., 
3 Aufl., 1891. 238; Miguhi, Syst. d. Bakt., 
2, 1900, 886.) From intestine of children. 

Pseudomonas weigmanni Migula. 
(Bakterie IV, Weigmann and Zirn, Cent, 
f. Bakt., 15, 1894, 466; Migula, Syst. d. 
Bakt., 2, 1900, 892.) From soapy milk. 

Pseudomonas zorkendorferi Migula. 
(Bacillus oogenes fluoresccns a, Zorken- 
dorfer. Arch. f. Hyg., 16, 1893, 392; 
Migula, Syst. d. Bakt., 2, 1900, 897.) 
From hens' eggs. 

Appendix II : The following polar flagel- 
late organism has been described from 
activated sludge. H. Winogradsky has 
also described polar flagellate forms from 
the same source that form zoogloea 
(Compt. rend. Acad. Sci. Paris, 200, 



1935, 1887; Ann. Inst. Pasteur, 58, 1937, 
333). 

Zoogloea ramigera Kruse emend. 
Butterfield. (Kruse, in Fliigge, Die 
Mikroorganismen, 3 Aufl., 1, 1896, 68; 
Butterfield, Public Health Reports, 50, 
1935, 671; Culture No. 50, Wattle, Pub. 
Health Reports, 57, 1942, 1519.) 

Rods: 1 by 2 to 4 microns, with 
rounded ends. Non-spore-forming. 
Capsules present. Motile with a single 
long polar flagellum. Gram-negative. 

Gelatin : No liquefaction. 

Grows better in aerated liquid media. 

Agar : Scant growth. 

Indole not formed. 

No HoS produced. 

No acid or gas from carbohydrates. 

Nitrites not produced from nitrates. 

Optimum pH 7.0 to 7.4. 

Optimum temperature 28° to 30 °C. 
Good growth at 20° and at 37 °C. Mini- 
mum temperature 4°C. 

Strict aerobe. 

Distinctive character : Oxidizes sewage. 

Source : Isolated from activated sludge. 

Habitat: Produces zoogloeal masses in 
activated sludge. 



Genus II. Xanthomonas Dowson* 

(Phytoinonas Bergey et al., Manual, 1st ed., 1923, 174 ; Dowson, Cent, f . Bakt., II Abt., 

100, 1939, 187.) 

Cells usually monotrichous, with yellow, water-insoluble pigment. Proteins are 
usually readily digested. Milk usually becomes alkaline. Hydrogen sulfide is pro- 
duced. Asparagin is not sufficient as an only source of carbon and nitrogen. Acid is 
produced from mono- and disaccharides. Mostly plant pathogens causing necrosis. 
From Gr. xanthus, yellow ; monas, a unit ; M. L. monad. 

The type species is Xanthomonas hyacinihi (Wakker) Dowson. 

Key to the species of genus Xanthomonas. 

1. Colonies yellow. 

a. Gelatin liquefied. 

b. Starch hydrolysis feeble. 

c. Nitrites not produced from nitrates. 

1. Xanthomonas hyacinthi. 

2. Xanthomonas pruni. 

3. Xanthomonas vitians. 



Prepared by Prof. Walter H. Burkhol der, Cornell Univ., Ithaca, N. Y., June, 1943. 



FAMILY PSEUDOMONADACEAE 151 

cc. Nitrites produced from nitrates. 

4. Xanthomonas beticola. 

5. Xanthomonas lactucae-scariolac. 

6. Xanthomonas ruhrilincans. 
bb. Starch hydrolysis strong. 

c. Nitrites not produced from nitrates. 

d. No brown pigment in beef-extract agar. 

7. Xanthomonas barbareae. 

8. Xanthomonas begoniac. 

9. Xanthomonas campestris. 

9a. Xanthomonas campestris var. armoraciae. 

10. Xanthomonas citri. 

11. Xanthomonas corylina. 

12. Xanthomonas cucurbitae. 

13. Xanthomonas dieffenbachiae. 

14. Xanthomonas holcicola. 

15. Xanthomonas incanae. 

16. Xanthomonas jitglandis. 

17. Xanthomonas lespedezae. 

18. Xanthomonas malvacearnm. 

19. Xanthomonas pelargonii. 

20. Xanthomonas phaseoli. 

20a. Xanthomonas phaseoli var. sojense. 

21. Xanthomonas plantaginis. 

22. Xanthomonas ricinicola. 

23a. Xanthomonas translucens f. sp. hordei. 
23b. Xanthomonas translucens i. sp. undidosa. 
23c. Xanthomonas translucens f. sp. secalis. 
23d. Xanthomonas translucens f. sp. hordei-avenae. 
23e. Xahthomonas translucens f. sp. cerealis. 

24. Xanthomonas vascular inn. 

25. Xanthomonas vesicatoria. 

25a. Xanthomonas vesicatoria var. raphani. 
dd. Brown pigment produced in beef -extract media. 

26. Xanthomonas nakalac. 

20b. Xanthomonas phaseoli var. fuscans. 
cc. Nitrites produced from nitrates. 

27. Xanthomonas papavericola. 
. ccc. Ammonia formed in nitrate media. 

28. Xanthomonas alfalfae. 
bbb. Starch not hydrolyzed. 

c. Nitrites produced from nitrates. 

29. Xanthomonas acernae. 

cc. Nitrites not produced from nitrates. 

30. Xanthomonas carotae. 

31. Xanthomonas hederae. 

32. Xanthomonas phormicola. 
25. Xanthomonas vesicatoria. 

ccc. Ammonia formed in nitrate media. 

33. Xanthomonas geranii. 



152 MANUAL OF DETERMINATIVE BACTERIOLOGY 

bbbb. Starch hydrolysis not reported. 

c. Nitrites produced from nitrates. 

34. Xanthomonas antirrhini. 

35. Xanthomonas heterocea. 
cc. Nitrites not produced from nitrates. 

36. Xanthomonas gummisudans . 

37. Xanthomonas lactxicae. 

38. Xanthomonas nigromaculans . 
aa. Gelatin not liquefied. 

b. Starch not hydrolyzed. 

39. Xanthomonas oryzae. 
aaa. Gelatin not reported. 

b. Starch hydrolyzed. 

40. Xanthomonas celebensis. 
2. Colonies whitish to cream. 

a. Gelatin liquefied. 

b. Starch hydrolyzed. 

c. Nitrites produced from nitrates. 

41. Xanthomonas panici. 

42. Xanthomonas proteomaculans . 

43. Xanthomonas majuhotis. 
cc. Nitrites not reported. 

44. Xanthomonas ruhrisnhalhicans . 
bb. Starch not reported. 

45. Xanthomonas cannae. 

46. Xanthomonas zingiberi. 

47. Xanthomonas conjaci. 

1. Xanthomonas hyacinthi (Wakker) Milk: Casein is precipitated and 

Dowson. {Bacterium hyacinthi Wakker, digested. Tyrosine crystals produced. 
Botan. Centralblatt, 14-, 1883, 315; Nitrites not produced from nitrates. 

Bacillus hyacinthi Trevisan, I generi e le Indole : Slight production, 

specie delle Batteriacee, 1889 ; 19 ; Pseudo- Hydrogen sulfide is produced. 

monas hyacinthi Erw. Smith, Bot. Acid, no gas, from glucose, fructose, 

Gazette, ^4) 1897, 188; Phyiomonas galactose, suci'ose and maltose. 
hyacinthi Bergey et al., Manual, 1st ed., Starch: Hydrolysis slight. 

1923, 177; Dowson, Cent. f. Bakt., II Optimum temperature 28° to 30°C. 

Abt., 100, 1939, 188.) From Gr. hy- Maximum 34° to 35 °C. Minimum4°C. 
acinthus, the hyacinth; M. L. Hi/a- Aerobic, with the exception of maltose, 

cinthus, a generic name. where it is facultative anaerobic. 

Description from Smitli, Div. Veg. Habitat: Produces a yellow rot of 

Phys. and Path., U. S. D. A. Bui. 26, hyacinth bulbs, Hyacinthus orientalis. 
1901, 40. 

Rods: 0.4 to 0.6 by 0.8 to 2 microns. 2. Xanthomonas pruni (Erw. Smith) 

Motile with a polar flagellum. Filaments Dowson. {Pseudomonas pruni Erw. 

present. Gram-negative. Smith, Science, N. S. 17, 1903, 456; 

Gelatin: Slow liquefaction. jBacieHfwi pr?<m Erw. Smith, Bacteria in 

Agar colonies: Circular, flat, moist, Relation to Plant Dis., 1, 1905, 171; 

shining, bright yellow. Media stained Bacillus pruni Holland, Jour. Bact., 5, 

brown. 1920, 220; Phytomonas pruni Bergey et 



FAMILY PSEUDOMONADACEAE 



153 



al., IManual, 1st ed., 1923, 179; Dowson, 
Cent. f. Bakt., II Abt., 100, 1939, 190.) 
From L. prunus, plum; M. L. Primus, a 
generic name. 

Probable synonym : Phytoinonas cerasi 
wraggi Sackett, Col. Agr. Exp. Sta. Rept., 
S8, 1925, 17; Pseudomonas cerasi wraggi, 
ibid.; Bacterium cerasi wraggi Elliott, 
Bact. Plant Pathogens, 1930, 111. 

Description from Dunegan, U. S. Dept. 
Agr., Tech. Bull. 273, 1932, 23. 

Rods : 0.2 to 0.4 by 0.8 to 1.0 microns. 
Capsules. Motile with a polar flagellum. 
Gram-negative. 

Gelatin: Liquefaction. 

Beef -extract agar colonies: Yellow, 
circular, smooth, convex, edges entire. 

Broth : Turbid becoming viscid. 

Milk : Precipitation of casein and 
digestion. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not produced. Hy- 
drogen sulfide produced (Burkholder). 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942, 600). 

Acid from arabinose, xylose, glucose, 
fructose, galactose, mannose, maltose, 
lactose, sucrose, raffinose, melezitose. 

Starch is hydrolyzed (slight). 

Aerobic. 

Optimum temperature 24° to 29 °C. 
Maximum 37 °C. 

Source : Smith isolated the pathogen 
from Japanese plums. 

Habitat : Pathogenic on plum {Prumis 
salicina), peach {P. persica), apricot 
(P. armenicca) , etc. 

3. Xanthomonas vitians (Brown) Starr 
and Weiss. (Bacterium vitians Brown, 
Jour. Agr. Res., 13, 1918, 379 ; Phytomonas 
vitians Bergey et al.. Manual, 1st ed., 
1923, 183; Pseudomonas vitians Stapp, 
in Sorauer, Handb. d. Pflanzenkrank., 
2, 5 Aufl., 1928, 287; Starr and Weiss, 
Phytopath., 33, 1943, 316.) From Latin, 
vitians, injuring, infecting. 

Rods: Motile with bipolar flagella. 
Gram-negative. 



Gelatin: Slow liquefaction. 

Beef -extract agar colonies: Circular, 
smooth, thin, cream to cream-yellow. 

Broth : Turbid with yellow ring. 

Milk: Clears and turns alkaline. 

Nitrites not produced from nitrates. 

Indole: Feeble production. 

Hydrogen sulfide : Feeble production. 

Acid but not gas from glucose. 

Starch : Feeble hydrolysis. 

Optimum temperature 26° to 28 °C. 
Maximum 35 °C. Minimum 0°C. 

Aerobic. 

Source : Isolated from the stem of 
diseased lettuce plants from South 
Carolina. 

Habitat : Pathogenic on lettuce, Lac- 
tvca sativa. 

4. Xanthomonas beticola (Smith, 
Brown and Townsend) comb. nov. {Bac- 
terium beticolum Smith, Brown and 
Townsend, U. S. Dept. Agr., Bur. Plant 
Ind., Bui. 213, 1911, 194; Pseudomonas 
beticola Holland, Jour. Bact., 5, 1920, 
224; Phytomonas beticola Bergey et al.. 
Manual, 1st ed., 1923, 182.) From Latin, 
beta, beet; -cola, dweller. 

Description from Brown, Jour. Agr. 
Res., 37, 192S, 167, where the species is 
referred to as Bacterium beticola (Smith, 
Brown and Townsend) Potebnia. 

Rods: 0.4 to 0.8 by 0.6 to 2.0 microns. 
Motile with 1 to 4 polar flagella. Cap- 
sules. Gram-variable. 

Gelatin: Liquefaction. 

Beef-agar slants : Moderate filiform 
growth, flat, glistening, yellow. 

Broth : Turbid, yellow ring, abundant 
sediment. 

Milk: Coagulation and peptonization. 

Indole not formed. 

Hydrogen sulfide formed. 

Nitrites produced from nitrates. 

Acid from glucose, sucrose, maltose, 
mannitol. No acid from lactose. 

Starch hydrolysis feeble. 

Optimum temperature 29 °C. Maxi- 
mum 39°C. Minimum 1.5°C. 



154 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Optimum pH 6.5. Maximum 9.0 to 
9.5. Minimum 4.5 to 4.8. 

Tolerates salt up to 9 per cent. 

Aerobic. 

Source : Isolated from galls on sugar 
beets collected in Colorado, Kansas, 
and Virginia. 

Habitat : Produces gall on sugar beets 
and on garden beets. 

Note : It is doubtful whether this 
species belongs in this genus. 

5. Xanthomonas lactucae-scariolae 

(Thornberry and Anderson) comb. nov. 
{Phytomonas lactucae-scariolae Thorn- 
berry and Anderson, Phytopath., 27, 
1937, 109.) From Lactuca scariola, the 
host. 

Rods: 0.5 to 1.0 by 1.0 to 1.5 microns. 
Motile with 1 or 2 polar flagella. Chains 
present. Capsules. Gram-negative. 

Gelatin : Slow liquefaction. 

Glucose agar colonies : Round, entire, 
finely granular, amber yellow. 

Broth: Turbid. No pellicle. A yel- 
low rim. 

Milk: Slight acid, and peptonization. 

Nitrites are produced from nitrates. 

Hydrogen sulfide not formed. 

No gas from carbon sources. 

Starch: Slight diastatic activity. 

Optimum temperature 25 °C. Maxi- 
mum 35°C. Minimum 7°C. 

Aerobic. 

Source : Isolated from necrotic lesions 
on wild lettuce. 

Habitat : Pathogenic on wild lettuce, 
Lactuca scariola, but not on cultivated 
lettuce, Lactuca sativa. 

6. Xanthomonas rubrilineans (Lee et 
al. ) Starr and Burkholder. {Phytomonas 
rubrilineans Lee, Purdy, Barnum and 
Martin, Haw^aiian Sugar Planters' Assoc. 
Bull., 1925, 25; Pseudomonas rubrilineans 
Stapp, in Sorauer, Handb. d. Pflanzen- 
krank., 2, 5 Aufi., 1928, 35; Bacterium 
rubrilineans Elliott, Man. Bact. Plant 
Path., 1930, 195; Starr and Burkholder, 
Phytopath., 32, 1942, 600.) From L. 
ruber, rsd; lineans, striping. 



Rods: 0.7 by 1.67 microns. Motile 
with 1 or seldom more polar flagella. 
Gram-negative. 

Gelatin: Liquefaction. 

Agar (Beef -extract + glucose) col- 
onies: Small, smooth, glistening, buff to 
yellow. 

Broth: Turbid with pellicle. Sedi- 
ment. 

Milk : Casein precipitated and digested. 

Nitrites are produced from nitrates. 

Indole not produced. 

Hydrogen sulfide not formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., S2, 1942, 600). 

Acid from glucose, fructose, arabinose, 
xylose, lactose, sucrose, raffinose and 
mannitol. 

Starch: Slight hydrolysis. 

Growth range, pH 5.4 to pH 7.3. 

Facultative anaerobe. 

Source : Description from 3 cultures 
isolated from the red stripe lesions in 
sugar cane. 

Habitat : Pathogenic on sugar cane. 

7. Xanthomonas barbareae Burkholdei , 
(Burkholder, Phytopath., 31, 1941, 348- 
Phytomonas barbareae Burkholder, ibid.) 
From M. L. Barbarea, a generic name. 

Rods : 0.4 to 0.95 by 1.0 to 3.15 microns. 
Motile with a single polar flagellum. 
Gram-negative. 

Gelatin : Liquefaction. 

Beef -extract peptone colonies : Cir- 
cular, yellow, smooth, butyrous, growth 
moderate. 

Potato glucose agar : Growth abun- 
dant, pale yellow. Mucoid. 

Broth : Turbid, yellow granular ring. 

Milk: Soft curd, with clearing and 
production of tyrosine crystals. Litmus 
reduced. 

Nitrates utilized but no nitrites formed. 
Asparagine and nitrites not utilized. 

Hydrogen sulfide produced. 

Indole not formed. 

Lipolytic (Starr and Burkholder, loc. 
cit.). 

Acid from glucose, galactose, xylose, 
maltose, sucrose, and glycerol. Alkali 



FAMILY PSEUDOMONADACEAE 



155 



produced from salts of malonic, citric, 
malic, and succinic acids. Rhamnose, 
salicin and hippuric acid salts not 
utilized. 

Starch hydrolyzed. 

Aerobic. 

Distinctive characters : Similar to 
Xanthomonas campestris but does not 
infect cabbage, cauliflower or horseradish. 

Source : From black rot of winter cress, 
Barbarea vulgaris. 

Habitat : Pathogenic on leaves and 
stems of Barbarea vulgaris. 

8. Xanthomonas begoniae (Takimoto) 
Dowson. {Bacterium begoniae Takimoto, 
Jour. Plant. Protect., 21, 1934, 262; 
Pseudomonas begoniae Stapp, Arbeiten 
Biol. Reichsanst. f. Land- und Forstw., 
22, 1938, 392; Phytomonas begoniae Burk- 
holder, in Manual, 5th ed., 1939, 162; 
Dowson, Cent. f. Bakt., II Abt., 100, 
1939, 190.) From M. L. Begonia, a 
generic name. 

Probable synonyms : Bacterium be- 
goniae Buchwald noni. nud., Gartner- 
Tidende, ^5, 1933, 1 ; Phytomonas flava 
begoniae Wieringa, Tidschr. Plantziekt., 
^1 , 1935, 312; Bacterium flavozonala 
McCulloch, Jour. Agr. Res., 54, 1937, 859 
{Xanthomonas flavozonatum Dowson, loc. 
cil.). 

Translated by Dr. K. Togashi. 

Rods: 0.5 to 0.6 by 1.2 to 2.0 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: No liquefaction. Liquefac- 
tion (Wieringa, loc. cit., McCulloch, 
loc. cit., Dowson, loc. cit., and Stapp, 
loc. cit.). 

Potato agar colonies: Circular, convex, 
smooth, moist, shining, yellow. 

Broth: Turbid. Yellow pellicle and 
precipitation. 

Milk : No coagulation. Casein digested 
Alkaline. 

Nitrites not produced from nitrates. 

Indole not produced. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942, 600). 



Starch hydrolyzed (Dowson, Jour. 
Roy. Hort. Soc, 63, 1938, 289). 

No acid or gas in peptone broth from 
glucose, sucrose, lactose or glycerol. 
Acid from glucose, sucrose, lactose, 
mannitol and glycerol in peptone-free 
medium (McCulloch, loc. cit.). 

Optimum temperature 27 °C. Maxi- 
mum 37°C. Minimum 1° to 3°C. 

Source : Isolated from leaf spot of 
begonia. 

Habitat : Pathogenic on Begonia spp. 

9. Xanthomonas campestris (Pam- 
mel) Dowson. {Bacillus campestris 
Pammel, Iowa Agr. Exp. Sta. Bull. 27, 
1895, 130; Pseudomonas campestris Erw. 
Smith, Cent. f. Bakt., II Abt., 3, 1897, 
284; Bacterium campestris (sic) Chester, 
Del. Col. Agr. Exp. Sta. Ann. Rept., 9, 
1897, 110; Phytomonas campestris Bergey 
et al., Manual, 1st ed., 1923, 176; Dowson, 
Cent. f. Bakt., II Abt., 100, 1939, 190.) 
From L. campestris of the field. 

Description from McCulloch (Jour. 
Agr. Res., 38, 1929, 278). Species is 
probably composed of several varieties. 
See descriptions by Mekta, Ann. Appl. 
Biol., 12, 1925, 330; Paine and Nirula, 
Ann. Appl. Biol., 15, 1928, 46; Wormald 
and Frampton, Ann. Rept. East. Mall. 
Res. Sta., 1926 and 1927, II Supplement, 
1928, 108; and others. 

Rods: 0.3 to 0.5 by 0.7 to 2.0 microns. 
Motile with a polar flagellum. Capsules. 
Gram-negative. 

Gelatin: Liquefied. 

Beef agar colonies : Wax yellow, round, 
smooth, shining, translucent, margins 
entire. 

Broth : Turbid with j'ellow rim and 
sometimes a pellicle. 

Milk: Casein digested with the forma- 
tion of tyrosine crystals. Alkaline. 

Nitrites not produced from nitrates. 

Indole formation weak. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942,600). 

Acid, no gas, from glucose, sucrose, 
lactose, glycerol and mannitol. 



156 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Starch is hydrolyzed. 

Optimum temperature 28° to 30 °C. 
Maximum 36 °C. 

Aerobic. 

Distinctive characters: Causes a vas- 
cular infection in cabbage, cauliflower 
and rutabagas. 

Source: Pammel {loc. cit.) first isolated 
the pathogen from diseased rutabagas. 

Habitat: Pathogenic on cabbage, cauli- 
flower and other related species. 

9a. Xanlhomonas carnpestris var. ar- 
moraciae (McCulloch) Burkholder. 
(Bacterium camjiestre var. armoraciae 
McCulloch, Jour. Agr. Res., S8, 1929, 
269; Phyiomonas carnpestris var. ar- 
moraciae Bergey et al., Manual, 3rd ed., 
1930, 251; Burkholder, Phytopath., 32, 
19-12, 601.) From Or. Armoracia, the 
horse radish. 

Cultural characters same as Xanlho- 
monas carnpestris. 

Distinctive characters : Causes a leaf 
spot of horse radish. No vascular in- 
fection. 

Source : Isolated from diseased horse- 
radish leaves collected in Washington, 
D. C, Virginia, Connecticut, Iowa and 
Missouri. 

Habitat : Pathogenic on horse radish 
and related species. 

10. Xanthomonas citri (Hasse) Dowson. 
(Pseudomonas citri Hasse, Jour. Agr. 
Res., 4, 1915, 97; Bacterium citri Doidge, 
Union So. .\frica, Dept. Agr. Sci. Bui. 8, 
1916, 20; Bacillus citri Holland, Jour. 
Bact., 5, 1920, 218; Phyiomonas citri 
Bergey et al., Manual, 1st ed., 1923, 181 ; 
Dowson, Cent. f. Bakt., II Abt., 100, 
1939, 190.) From M. L. Citrus, a generic 
name. 

Rods: 0.5 to 0.75 by 1.5 to 2 microns, 
occurring in chains. Motile with a single 
polar flagellum. Gram-negative. 

Gelatin: Liquefied. 

Beef agar colonies : Appear in 36-48 
hours, circular, smooth, raised, dull 
yellow. 

Broth: Turbid in 24 hours. A yellow 
ring formed. 



Milk : Casein is precipitated. 

Nitrites not produced from nitrates. 

Hydrogen sulfide produced (Reid, New 
Zealand Jour. Sci . and Tech. , .2^, 1938, 60 ) . 

Indole not formed. 

No gas from glucose, lactose or man- 
nitol. 

Starch h3'drolyzed (Reid, loc. cit.). 

Aerobic. 

Optimum temperature, 25° to 34 °C. 
Maximum 38°C. Minimum 10°C. 
(Okabe, Jour. Soc. Trop. Agr., 4, 1932, 
476). 

Source : Isolated from canker on orange. 

Habitat: Produces a canker on many 
species of Citrus and related plants. 

11. Xanthomonas corylina Miller, Bol- 
len, Simmons, Gross, and Barss. (Miller 
et al., Phytopath., 30, 1940, 731; Phyio- 
monas corylina Miller et al., ibid.) 
From Gr. corylus, the hazelnut; M. L. 
corylina, of the hazel nut. 

Rods: 0.5 to 0.7 by 1.1 to 3.8 microns. 
Motile with a polar flagellum. Capsules. 
Gram-negative. 

Gelatin : Liquefaction. 

Nutrient glucose-agar streaks : Abun- 
dant growth, filiform, convex, glistening, 
smooth, opaque, pale lemon yellow, viscid. 

Broth: Turbid. Ring formed in 2-5 
days. 

Milk : Enzymatic curd that is slowly 
digested. Litmus reduced. Crystal for- 
mation (Burkholder). 

Nitrites not produced from nitrates. 

Nitrogen sources utilized are peptone, 
aspartic acid, alanine, leucine, sodium 
ammonium phosphate, allantoin, tyro- 
sine, uric acid and brucine. 

Indole is not produced. 

Hydrogen sulfide not produced on lead 
acetate agar. H2S produced after Zobell 
and Feltham's method (Burkholder). 

Selenium dioxide reduced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., S;g, 1942, 600). 

Acid, no gas from glucose, fructose, 
galactose, lactose, sucrose, maltose, xy- 
lose, raffinose, mannitol, glycerol, and 
starch. Alkali from salts of citric, lactic, 



FAMILY PSEUDOMONADACEAE 



157 



malic and succinic acids. Arabinose, 
rhamnose, dulcitol, salicin, inulin, and 
cellulose not utilized. 

Starch is hydrolyzed. 

Optimum temperature 28° to 32°C. 
Maximum 37 °C. Minimum 5° to 7°C. 
Thermal death point 53° to 55 °C. 

pH range for growth: pH 5.2 to 10.5. 
Optimum pH 6 to 8. 

Strict aerobe. 

Distinctive characters : Cultural char- 
acters the same or similar to Xantho- 
monas juglandis. The two species do 
not cross-infect. 

Source : 26 isolates from widely scat- 
tered filbert orchards in Oregon and 
Washington. 

Habitat: Pathogenic on filberts (Cory' 
lus avellana and C. maxima). 

12. Xanthomonas cucurbitae (Bryan) 
Dowson. {Bacterium cucurbitae Bryan, 
Science, 63, 1926, 165; Bryan, Jour. Agr. 
Res., 40, 1930, 389; Phytomonas cucur- 
bitae Bergey et al., Manual, 3rd ed., 
1930, 251 ; Pseudomonas cucurbitae Stapp, 
Bot. Rev., 1, 1935, 408; Dowson, Cent. f. 
Bakt., II Abt., 100, 1939, 190.) From L. 
curczirbita, a gourd; M. L. Cucurbita, a 
generic name. 

Rods : 0.45 to 0.6 by 0.5 to 1.3 microns. 
Motile, usually with a single polar flagel- 
lum. Gram-negative. 

Gelatin: Liquefied. 

Beef-agar slants: Growth moderate, 
mustard yellow, undulating margins, 
viscid to butyrous. 

Broth: Moderately turbid. Ring and 
yellow sediment. 

Milk : Precipitation of casein and diges- 
tion. Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced. 

Acid from glucose, galactose, fructo.se, 
lactose, maltose, sucrose and glycerol. 
No acid from mannitol. 

Starch is hydrolyzed. 

Optimum temperature 25° to 30 °C. 
Maximum 35 °C. 



Optimum pH 6.5 to 7.0. Limits of 
growth pH 5.8 to 9.0. 

Slight growth in 5 per cent salt. 

Aerobic. 

Source : Species first isolated from 
squash. 

Habitat: Causes a leaf spot of squash 
and related plants. 

13. Xanthomonas dieffenbachiae (Mc- 
Culloch and Pironc) Dowson. {Phy- 
tomonas dieffenbachiae McCulloch and 
Pirone, Phytopath., 29, 1939, 962; Bac- 
terium dieffenbachiae McCulloch and 
Pirone, ibid.; Dowson, Trans. Brit. 
Mycol. Soc, 26, 1943, 12.) From M. L. 
Dieffenbachia, a generic name. 

Rods: 0.3 to 0.4 by 1.0 to 1.5 microns. 
Motile with a single polar flagellum. 
Capsules. Gram-negative. 

Gelatin: Liquefied. 

Beef-infusion peptone agar colonies: 
Slow growing, circular, flat, smooth, 
translucent. Butyrous. Massicot to 
Naples yellow. 

Broth: Turbid. Yellow rim or slight 
pellicle. 

Milk : Slow peptonization and forma- 
tion of tyrosine crj^stals. Litmus re- 
duced. 

Nitrites not formed from nitrateaft 

Indole not produced. 

Hydrogen sulfide produced. 

Acid from glucose, sucrose, lactose, 
galactose, fructose and glycerol . Growth 
but no acid in maltose and mannitol. 

Starch moderately hydrolyzed. 

Optimum temperature 30° to 31 °C. 
Maximum 37° to 38 °C. Minimum 5°C. 

Aerobe. 

Source : Seven isolates from diseased 
leaves of Dieffenbachia picta. 

Habitat : Pathogenic on Dieffenbachia 
picta. Artificial infection of Dracaena 
fragrans . 

14. Xanthomonas holcicola (Elliott) 
Starr and Burkholder. {Bacterium hol- 
cicola Elliott, Jour. Agr. Res., 40, 1930. 
972; Phytomonas holcicola Bergey et al.. 
Manual. 4th ed., 1934, 271; Pseudomonas 



158 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



holcicola Stapp, Bot. Rev., 1, 1935, 407; 
Starr and Burkholder, Phytopath., S2, 
1942, 600.) From Gr. holcus, sorghum, 
-cola, dweller; M. L. Holcus, a generic 
name. 

Rods: 0.75 by 1.58 microns. Motile 
with 1 or 2 polar flagella. Capsules. 
Gram-negative. 

Gelatin: Liquefied. 

Beef-infusion peptone agar colonies : 
Round, umbonate, glistening, smooth, 
translucent to opaque, wax yellow, buty- 
rous. 

Broth: Trace of growth in 24 hours. 
Later turbid with a slight ring. 

Milk: Casein precipitated and pep- 
tonized. Alkaline. 

Nitrite production doubtful. 

Indole not produced. 

Hydrogen sulfide is produced. 

Lipolytic (Starr and Burkholder, loc. 
cit.). 

Acid, no gas, from sucrose. 

Starch is hj'drolyzed. 

Optimum temperature 28° to 30°C. 
Maximum 36° to 37°C. Minimum 4°C. 

Optimum pH 7.0 to 7.5. Growth 
range pH 5.5 to 9.0. 

Source : Isolated from many collections 
of sorghum leaves showing a streak 
disease. 

Habitat: Pathogenic on leaves of 
Holcus sorghum and //. halepensis. 

15. Xanthomonas incanae (Kendrick 
and Baker) Starr and Weiss. (Phyto- 
monas incanae Kendrick and Baker, 
California Bull. 665, 1942, 10; Starr and 
Weiss, Phytopath., S3, 1943, 316.) From 
its host plant Malthiola incana; L. 
incanus, quite gray or hoary. 

Rods: 0.4 to 0.8 by 0.6 to 2.5 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin : Liquefied. 

Beef extract agar colonies : Round, 
smooth, convex or pulvinate, glistening, 
margin entire, picric yellow to amber color. 

Broth: Turbid. 

Milk : No coagulation. A clearing of 
the medium. 



Nitrites not produced from nitrates. 

Indole not formed. 

Lipolj^tic (Starr and Burkholder, Phy- 
topath., 32, 1942, 600). 

Acid but no gas from glucose, lactose, 
sucrose, mannitol, d-galactose, xj'lose, 
d-mannose, raffinose, trehalose, and glyc- 
erol. No acid from maltose, 1-arabinose, 
or rhamnose. 

Starch not hydrolyzed. Starch hy- 
drolyzed (Burkholder). 

Tolerates 3 per cent salt. 

Growth in beef broth at pH 4.4. 

Aerobic. 

Distinctive characters: Causes a dis- 
ease of flowering stock but not of cabbage. 
Differs from Xanthomonas campestris in 
that it does not utilize 1-arabinose, nor 
maltose. 

Source : Four isolates from diseased 
plants of Matlhiola incana. 

Habitat : Pathogenic on flowering 
stocks. 

16. Xanthomonas juglandis (Pierce) 
Dowson. (Psetidomonas juglandis Pierce, 
Bot. Gaz., 31, 1901, 272; Bacterium 
juglandis Erw. Smith, Bacteria in Rela- 
tion to Plant Dis., 1, 1905, 171; Bacillus 
juglandis Holland, Jour. Bact., 5, 1920, 
218; Phytomonas juglandis Bergey et al., 
Manual, 3rd ed., 1930, 247; Dowson, 
Cent. f. Bakt., II Abt., 100, 1939, 190.) 
From L. juglans (-andis), the walnut; 
M. L. Juglans, a generic name. 

Description taken from Miller et al., 
Phytopath., 30, 1940, 731. 

Rods: 0.5 to 0.7 by 1.1 to 3.8 microns. 
Motile with a polar flagellum. Capsules. 
Gram-negative. 

Gelatin: Liquefaction. 

Nutrient glucose-agar streaks : Abun- 
dant growth, filiform, convex, glistening, 
smooth, opaque, pale lemon yellow, 
viscid. 

Broth: Turbid. Ring formed in 2 to 
5 days. 

Milk : Enzymatic curd that is slowly 
digested. Litmus reduced. Crystal for- 
mation (Burkholder). 

Nitrites not produced from nitrates. 



FAMILY PSEUDOMONADACEAE 



159 



Nitrogen sources utilized are peptone, 
aspartic acid, alanine, leucine, sodium 
ammonium phosphate, allantoin, tyrosine 
uric acid and brucine. 

Indole is not produced. 

Hydrogen sulfide not produced on lead 
acetate agar. H2S produced after Zobell 
and Feltham's method (Burkholder). 

Selenium dioxide reduced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942, 600). 

Acid, no gas from glucose, fructose, 
galactose, lactose, sucrose, maltose, xy- 
lose, raffinose, mannitol, glycerol, and 
starch. Alkali from salts of citric, lactic, 
malic and succinic acid. Arabinose, 
rhamnose, dulcitol, salicin, inulin, and 
cellulose not utilized. 

Starch is hydrolyzed. 

Optimum temperature 28° to 32 °C. 
Maximum 37 °C. Minimum 5° to 7°C. 
Thermal death point 53° to 55°C. 

pH range for growth pH 5.2 to 10.5. 
Optimum pH 6 to 8. 

Source : Isolated from black spots on 
the leaves and nuts of English walnuts, 
Juglans regia. 

Habitat : Pathogenic on the walnut, 
Juglans spp. 

17. Xanthomonas lespedezae (Ayers, 
Lefebvre and Johnson) comh. nov. (Phij- 
tomonas lespedezae Ayers, Lefebvre and 
Johnson, U. S. Dept. Agr. Tech. Bui. 
704, 1939, 19.) From AI. L. Lespedeza, 
a generic name. 

Rods: 0.56 by 1.62 microns. Single, 
in pairs or occasional short chain. Mo- 
tile with 1 polar flagellum. Gram- 
negative. Capsules. 

Gelatin: Liquefied. Also egg albumin 
and blood serum. 

Nutrient agar colonies : Circular, 
raised, glistening, translucent, and vis- 
cid. Yellow. 

Broth : Turbid in 48 hours. 

Milk : Peptonized and becomes alka- 
line. 

Nitrites not produced from nitrates. 

Indole is produced after 11 days. 

Hydrogen sulfide is produced. 



No gas in carbohydrates. 

Starch hydrolyzed. 

Aerobic. 

Optimum temperature near 35 °C. 
No growth at 5°C. or at 40 °C. 

Source : Isolated from diseased Les- 
pedeza spp. collected in Virginia, New 
York and Illinois. 

Habitat : Pathogenic on Lespedeza spp. 

18. Xanthomonas malvacearum (Erw. 
Smith) Dowson. (Pseudomonas mal- 
vacearum Erw. Smith, U. S. Dept. Agr., 
Div. Veg. Phys. and Path., Bui. 28, 1901, 
153; Bacterium malvacearum Erw. Smith, 
Bact. in Rel. to Plant Diseases, 1, 1905, 
171 ; Bacillus malvacearum Holland, 
Jour. Bact., 5, 1920, 219; Phytomonas 
malvacearum Bergey et al.. Manual, 1st 
ed., 1923, 178; Dowson, Cent. f. Bakt., 
II Abt., 100, 1939, 190.) From L. 
malva, mallows; M. L. Malvaceae, a 
family name. 

Description from Elliott, Man. Bact. 
Plant Pathogens, 1930, 153; and Lewis, 
Phytopath., 20, 1930, 723. 

Rods : Motile with one polar flagellum. 
Gram-negative. 

Gelatin: Liquefaction. 

Agar slants : IVIoderate growth, convex, 
smooth, glistening, pale yellow, wavy to 
irregular margins. 

Broth : Slight to moderate turbidity. 
Sediment. 

Milk : Casein precipitated and slowly 
digested. 

Nitrites not produced from nitrates. 

Hydrogen sulfide is produced (Burk- 
holder). 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 600). 

Acid but not gas from glucose, galac- 
tose, fructose, xylose, lactose, maltose, 
sucrose, raffinose, glycerol, inulin and 
glycogen. Alkaline reaction from salts 
of acetic, citric, lactic and succinic acids. 
No fermentation of arabinose, mannitol, 
dulcitol, salicin, and salts of formic, 
o.xalic and tartaric acids (Lewis, loc. cit). 

Starch hydrolyzed (Lewis, loc. cit.). 



160 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Optimum temperature 25° to 30 °C. 
Maximum 36° to 38 °C. (Elliott, loc. cit.). 

Source : Isolated from angular leaf spot 
of cotton. 

Habitat : Pathogenic on cotton where - 
ever it is grown, causing a leaf spot, a 
stem lesion and a boll lesion. 

19. Xanthomonas pelargonii (Brown) 
Starr and Burkholder. {Bacterium pe- 
largoni Brown, Jour. Agr. Res., 28, 1923, 
372; Pseudomonas -pelargoni Stapp, in 
Sorauer, Handb. d. Pflanzenkrank, 2, 5 
Aufl., 1928, 181; Phytonionas pelargonii 
Bergey et al., Manual, 3rd ed., 1930, 
250; Starr and Burkholder, Phytopath., 
32, 1942, 600.) P'rom Greek, pelargus, 
the stork; M. L. Pelargonium, a generic 
name for the stork's bill geranium. 

Rods : .67 by 1 .02 mi crons . Capsules . 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: Slow liquefaction. 

Beef -agar colonies : Cream -colored, 
glistening, round, with delicate internal 
markings. 

Broth: Turbid in 24 hours. Incom- 
plete pellicle. 

Milk: Alkaline. Clearing in bands. 

Nitrites not produced from nitrates. 

Indole formation slight. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., S;2, 1942, 600). 

Slight acid but not gas from glucose, 
sucrose and glycerol. 

Starch hydrolysis feebly positive. 

Optimum temperature 27 °C. Maxi- 
mum 35 °C. 

No growth in broth plus 3.5 per cent 
salt. 

Aerobic. 

Source: Isolated from spots on leaves 
of Pelargonium from District of Colum- 
bia, Maryland and New Jersey. 

Habitat: Pathogenic on Pelargonium 
spp. and Geranium spp. 

20. Xanthomonas phaseoli (Erw. 
Smith) Dowson. {Bacillus phaseoli Erw. 
Smith, Bot. Gaz., 2^, 1897, 192; A. A. A. 



S. Proc, JiB, 1898, 288; Pseudomonas 
phaseoli Erw. Smith, U. S. Dept. Agr., 
Div. Veg. Phys. and Path., Bui. 28, 1901, 
1 ; Bacterium phaseoli Erw. Smith, 
Bact. in Rel. to Plant Dis., 1, 1905, 72; 
Phytonionas phaseoli Bergey et al.. 
Manual, 1st ed., 1923, 177; Dowson, 
Cent. f. Bakt., II Abt., 100, 1939, 190.) 
From Gr. phaseolus, the bean; M. L. 
Phaseolus, a generic name. 

Description from Burkholder, Cornell 
Agr. Exp. Sta. Mem. 127, 1930, 18; 
and Phytopath., 22, 1932, 609. 

Rods: 0.87 by 1.9 microns. Motile 
with a polar flagellum. Gram-negative. 

Gelatin : Liquefaction. 

Beef -extract agar colonies: Circular, 
amber yellow, smooth, butyrous, edges 
entire. 

Broth : Turbid in 24 hours. Yellow 
ring. 

Milk : Casein precipitated and digested. 
Alkaline. Tyrosine crystals formed. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942,600). 

Acid but not gas from glucose, galac- 
tose, fructose, arabinose, xylose, maltose, 
lactose, sucrose, raffinose and glycerol. 
Alkaline reaction from salts of acetic, 
malic, citric and succinic acids. Man- 
nitol, dulcitol, salicin and formic and 
tartaric acids not fermented. 

Starch is hydrolyzed. 

Aerobic. 

Very slight growth in beef broth plus 
4 per cent salt (Hedges, Jour. Agr. Res., 
^,9,1924,243). 

Distinctive character: Similar in cul- 
ture to Xanthomonas campestris, X. 
juglandis, X. vesicatoria, etc., but they 
do not cross infect . 

Habitat: Pathogenic on the bean 
{Phaseolus vulgaris), the hyacinth bean 
{Dolichos lablab), the lupine {Lupinus 
polyphillus) , etc. Not pathogenic on 
the soy bean {Glycine sp.), nor cowpea 
{Vigna sp.). 



FAMILY PSEUDOMONADACEAE 



161 



20a. Xanthomonasphaseoli var. sojensis 
(Hedges) Starr and Burkholder. {Bac 
terium phaseoli var. sojense Hedges 
Science, 56, 1922, 11; Jour. Agr. Res. 
^9, 1924, 229; Phytomonas phaseoli var 
sojense Burkholder, Phytopath., 20 
1930, 7; Starr and Burkholder, Phyto- 
path., 32, 1942, 600.) From M. L. the 
soy bean, Soja, a generic name; M. L. 
sojensis, of the soybean. 

Synonyms : Pseudomonas glycines Na- 
kano. Jour. Plant Protect. Tokyo, 6, 1919, 
39 {Bacterium glycines Elliott, Manual 
Bact. Plant Path., 1928, 133; Phytomonas 
glycines IVIagrou, in Hauduroy et al., 
Diet. d. Bact. Path., 1937, 358). (See 
Takimoto, Jour. Plant Protect. Tokyo, 
18, 1931, 29; and Okabe, Jour. Trop. 
Agr. Formosa, 4, 1932, 473.) 

Distinctive character: Differs from 
Xanthomonas phaseoli in that it infects 
the soy bean, Glycine max. 

Source : Isolated from pustules on the 
leaves and pods of soy bean, both in 
America and in Japan. 

Habitat : Pathogenic on the soy bean. 
Glycine max and the common bean, 
Phaseolus vulgaris. 

20b. Xanthomonas phaseoli var. fus- 
cans (Burkholder) Starr and Burkholder. 
{Phytomonas phaseoli vslt. fuscans Burk- 
holder, Cornell Agr. Exp. Sta. Mem. 126, 
1930, 22; Phytopath., 22, 1932, 699; 
Bacterium phaseoli var. fuscans Okabe, 
Jour. Soc. Trop. Agr. Formosa, 5, 1933, 
161 ; Pseudomonas phaseoli var. fuscans 
Stapp, Bot. Rev., 1, 1935, 407; Starr and 
Burkholder, Phytopath., 32, 1942, 600.) 
From L. fuscans, producing a brown 
color. 

Distinctive characters: Differs from 
Xantlwmonas phaseoli in that it produces 
a deep brown color in beef-extract-pep- 
tone media and in tyrosine media. 
Action on maltose negative or feeble. 

Source : Two cultures isolated ; one 
from a diseased bean leaf (1924) and a 
diseased pod (1927) collected in Switz- 
erland. 



Habitat: Pathogenic on beans, Phase- 
olus vulgaris, and related plants. 

21. Xanthomonas plantaginis (Thorn- 
berry and Anderson) comb. nov. {Phy- 
tomonas plantaginis Thornberry and 
Anderson, Phytopath., 27, 1937, 947.) 
From Latin, Plantago (-agrzms), plantain; 
1\I. L. Plantago, a generic name. 

Rods: 0.6 to 1.0 by 1.0 to 1.8 microns. 
Occurring singly or in chains. Capsules. 
Motile with 1 to 2 polar flagella. Gram- 
negative. 

Gelatin : Slight liquefaction. 

Glucose agar slant : Growth moderate, 
filiform, raised, opaque, yellow and 
A-iscid. 

Broth: Moderately turbid with ring. 

Milk: Slight acidity, nt) reduction of 
litmus. Peptonization. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide not produced. 

No appreciable amount of gas from 
carbohydrates. 

Starch is hydrolyzed. 

Optimum temperature 25°C. Mini- 
mum 12°C. Maximum 35°C. Thermal 
death point 50°C. 

Aerobic. 

Source : From diseased leaves of Plan- 
tago lanceolata in Illinois. 

Habitat: Pathogenic on Plantago spp. 

22. Xanthomonas ricinicola (Elliott) 
Dowson. {Bacterium ricini Yoshi and 
Takimoto, Jour. Plant Protect. Tokyo, 
15, 1928, 12; Bacterium ricinicola Elliott, 
Man. Bact. Plant Path., 1930, 193; Phijto- 
monas ricinicola Burkholder, in Manual, 
5th ed., 1939, 152; Dowson, Cent. f. 
Bakt., II Abt., 100, 1939, 190; Xantho- 
monas ricini Dowson, ibid.) From L. 
living on the castor bean ; M. L. Ricinus, 
a generic name. 

Rods: 0.4 to 0.9 by 1.3 to 2.6 microns. 
Capsules. Short chains. Motile with 
polar flagella. Gram-negative. 

Gelatin : Liquefaction. 

Nutrient agar colonies : Lemon yellow, 
changing to brown. 



162 



MANUAL OF DETERMINATI\'E BACTERIOLOGY 



Milk: Slightly acid. No coagulation. 
Peptonization. 

Nitrites not produced from nitrates. 

Acid but not gas from lactose. 

Starch hydrolyzed. 

Optimum temperature 29° to 30 °C. 
Maximum 39°C. Minimum 2.5°C. 

Aerobic. 

Source : Isolated from leaf -spot of 
castor-bean. 

Habitat : Pathogenic on Ricinus com- 
munis. 

23a. Xanthomonas transtucens f. sp. 
hordei Hagborg. (Canadian Jour, of 
Res., W, 1942, 317.) From L. trans- 
lucens, shining through, translucent, 
referring to the character of the lesion 
produced by this pathogen. Form name 
from Hordeum, a generic name. 

Synonyms : Bacterium iranslucens 
Jones, Johnson and Reddy, Jour. Agr. 
Res., 11, 1917, 637; Pseudomonas irans- 
lucens, ibid. ; Phytomonas iranslucens 
Bergey et al. Manual, 3rd ed., 1930, 252; 
Xanthomonas iranslucens Dowson, Cent, 
f. Bakt., II Abt., 100, 1939, 190. 

Rods: 0.5 to 0.8 by 1 to 2.5 microns. 
Motile with a single polar flagellum. 
Gram-negative. 

Gelatin: Liquefaction. 

Beef-peptone agar colonies : Round, 
smooth, shining, amorphous except for 
inconspicuous somewhat irregular con- 
centric striations within, wax-yellow 
tinged with old gold; margin entire. 

Broth : Turbidity becomes rather 
strong. Pellicle. 

Milk: Soft coagulum and digestion. 
Milk clears. Tyrosine crystals produced. 

Nitrites not produced from nitrates. 

Indole: Slight formation. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942,600). 

Ammonia from peptone. 

Acid but not gas from glucose, d- 
fructose, d-mannose, d-galactose, sucrose, 
lactose, and sometimes salicin. No uti- 
lization of 1-rhamnose, inositol, maltose, 



raffinose, inulin, d-mannitol, and dul- 
citol. 

Starch hydrolyzed. 

Optimum temperature 26 °C. Maxi- 
mum 36 °C. Minimum 6°C. 

Aerobic. 

Distinctive characters : All forms of 
Xanthomonas iranslucens have the same 
cultural characters. They differ mainly 
in pathogenicity. This form is patho- 
genic on barley, Hordeum spp. ; but not 
on oats, Avena spp., rye, Secale cereale 
nor on wheat, Triiicum spp. 

Source : Isolated from leaves and seed 
of barley, Hordeum vulgare. 

Habitat : Occurs naturally on barley. 

23b. Xanthomonas iranslucens f. sp. 
undulosa (Smith, Jones and Reddy) Hag- 
borg. {Bacterium iranslucens var. undu- 
losum Smith, Jones and Reddy, Science, 
50, 1919, 48; Pseudomonas iranslucens 
var. undulosa Stapp, in Sorauer, Handb. 
d. Pflanzenk., 2, 5 Auf., 1928, 17; Phyto- 
monas iranslucens var. undulosa Hagborg, 
Canadian Jour. Res., H, 1936, 347; Hag- 
borg, Canadian Jour. Res., 20, 1942, 317.) 
From L. unda; M.L. undulosus , undulate, 
referring to the undulation of the colony. 

Distinctive characters : Cultural char- 
acters same as all forms of Xanthomonas 
iranslucens. Pathogenic on wheat, Tri- 
ticium spp., barley, Hordeum spp. and 
rye, Secale cereale but not on oats, 
Avena spp. 

Source : Isolated repeatedly from black 
chaff of wheat. 

Habitat : Usually found on wheat caus- 
ing the black chaff, and on rye. 

23c. Xanthomonas iranslucens f. sp. 
secalis (Reddy, Godkin and Johnson) 
Hagborg. {Bacterium iranslucens var. 
secalis Reddy, Godkin and .lohnson. 
Jour. Agr. Res., 28, 1924, 1039; Psew- 
domona iranslucens var. secalis Stapp, 
in Sorauer, Handb. d. Pflanzenkr., 2, 5 
Aufl., 1928,24; Phytomonas iranslucens 
var. secalis Burkholder, in Manual, 5th 
ed., 1939, 160;Hagborg, Canadian Jour. 



FAMILY PSEUDOMONADACEAE 



163 



Res., 20, 1942, 317.) From M.L. 5eca/e, 
a generic name. Distinctive characters: 
Cultural characters same as other forms 
of Xantliomonas translucens. This form 
pathogenic on rye Secale cereale, but 
not on Triticnm spp., Hordeum spp. 
nor Avena spp. 

Source : Isolated from leaf spot on rye, 
Secale cereale. 

Habitat : Pathogenic on rye. 

23d. Xanthomonas translucens f. sp. 
hordei-avenae Hagborg. (Canadian Jour. 
Res. JO, 1942,317.) FromM.L. Hordeum 
and Avena, generic names. 

Distinctive characters : Cultural char- 
acters same as other forms of Xantho- 
monas translucens . Pathogenic on barley, 
Hordeum spp. and oats, Avena spp., but 
not on wheat, Triticium spp., nor rye, 
Secale cereale. 

Source : Isolated 6 times from barley 
at various places in Canada. 

Habitat : Occurs naturally on barley. 

23e. Xanthomonas translucens i. sp. 
cerealis Hagborg. (Canadian Jour. Res., 
20, 1942, 317.) From L., of cereal. 

Distinctive characters : Cultural char- 
acters same as other forms of Xantho- 
monas translucens . Pathogenic on wheat, 
Triticum spp.; oats, Avena spp.; barley, 
Hordeum spp. ; and rj^e, Secale cereale. 

Source : Isolated from wheat in Canada. 

Habitat: Occurs naturallj^ on wheat. 

24. Xanthomonas vasculorum (Cobb) 
Dowson. {Bacillus rasciilarum (sic) 
Cobb, Agr. Gaz. of New South Wales, 4, 
1S93, 777 ; Abst. in Cent, f . Bakt., II Abt., 
/, 1895,41 ; Bacterium vascxdarum Migula, 
Syst. d. Bakt., 2, 1900, 512; Pseudomonas 
vascularinn Erw. Smith, U. S. Dept. Agr., 
Div. Veg. Phys. and Path., Bui. 28, 1901, 
153; Phytomonas vascularum Bergey et 
al.. Manual, 1st ed., 1923, 179; Dowson, 
Cent. f. Bakt., II Abt., 100, 1939, 190.) 
From L. vasculum, a small vessel ;M. L. 
the vascular system. 

Note: Erw. Smith (Bact. in Rel. to 



Plant Dis., 3, 1914, 88) states that prob- 
ably Spegazzini (El Polville de la Cana 
de Azucar, June, 1895, La Plata, Supl. 
Rev. Azuc, Buenos Aires, No. 16, 1895) 
reported the disease caused by Xantho- 
monas vasculorum but that Bacillus 
sacchari Spegazzini which he claimed 
to be the pathogen, was a saprophyte. 

Description from Smith (loc. cit., 54). 

Rods : 0.4 by 1.0 microns. Motile with 
a polar flagellum. Gram-variable. 

Gelatin : Liquefaction feeble. Lique- 
faction good (Burkholder). 

Beef -extract agar colonies : Pale j^ellow,. 
smooth, glistening, not noticeably viscid. 

Broth: Good growth. 

Milk: Alkaline. 

Nitrites not produced from nitrates. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 82, 1942, 600). 

Acid but not gas from glucose, fruc- 
tose andglj'cerol. 

Starch hydrolyzed (Burkholder). 

Optimum temperature 30°C. Maxi- 
mum 35° to 37.5°C (Elliott, loc. cit.). 

Habitat : Pathogenic on sugar cane, 
Saccharum officinarum, causing a bac- 
terial gummosis. 

25. Xanthomonas vesicatoria (Doidge) 
Dowson. {Bacterium vcsicatorium 
Doidge, Jour. Dept. Agr,, S. Africa, 1, 
1920, 718; also Ann. Appl. Biol., 7, 1921, 
428; Pseudomonas vesicatoria Stapp, in 
Sorauer, Handb. d. Pflanzenkrank., 2, 
5 Aufi., 1928, 259; Phytomonas vesica- 
toria Bergey et al., Manual, 3rd ed., 1930, 
253; Dowson, Cent. f. Bakt., II Abt., 
100, 1939, 190. ) From L. vesica, a blister ; 
M. L. vesicatorius, causing blisters. 

Synonyms : Gardner and Kendrick 
(Phytopath., 13, 1923, 307) list Pseudo- 
monas exitiosa Gardner and Kendrick 
(Phytopath., 11, 1921, 55; Bacterium 
exitiosum Gardner and Kendrick, Jour. 
Agr. Res., 21, 1921, 141; Phytomonas 
exitiosa Bergey et al., Manual, 1st ed., 
1923, 183) and an unnamed species, 
Higgins (Phytopath, 12, 1922, 513). 

Rods : 0.6 to 0.7 by 1.0 to 1.5 microns 



164 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Motile with a polar flagellum. Capsules. 
Gram-positive. Gram-negative (Gard- 
ner and Kendrick; and Higgins). 

Gelatin : Liquefaction. 

Nutrient agar colonies: Good growth. 
Circular, wet-shining, Naples yellow, 
edges entire. 

Milk: Casein precipitated and slowly 
digested. Tyrosine crystals. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced (Burk- 
b >ider). 

Lipolytic (Starr and Burkholder, Phy- 
topath.,3;?, 1942, 600). 

Acid but not gas from glucose, fruc- 
tose, sucrose, lactose, galactose, glycerol 
and dextrin. 

Certain strains hydrolyze starch, others 
do not (Burkholder and Li, Phytopath., 
31, 1941, 753). 

Optimum temperature 30°C. 

Source : Isolated from spotted tomato 
fruits in South Africa. 

Habitat : Pathogen on tomatoes, Lycu- 
persicon esculentum and peppers, Capsi- 
cum annuum. 

25a. Xanthomonas vesicatoria var. 
raphani (White) Starr and Burkholder. 
{Bacterium vesicatoria var. raphani 
White, Phytopath., 20, 1930, 653; Phylo- 
monas vesicatoria var. raphani Burk- 
holder, in Manual, 5th ed., 1939, 154; 
Starr and Burkholder, Phytopath., 32, 
1942, 600.) From M. L. Raphanus, the 
radish, a generic name. 

Distinctive characters : Cultural char- 
acters similar to Xanthomonas vesica- 
toria, but differs in that it is able to attack 
radishes, turnips, and other crucifers. 
Differs from Xanthomonas campestris in 
that it does not cause a vascular disease, 
and differs from Xanthomonas cainpestris 
var. armoraciae in that it is not patho- 
genic on horseradish. 

Source : Isolated from leaf spots of 
radish and turnips in Indiana. 

Habitat : Pathogenic on radish, turnips, 
and other crucifers ; and on tomato and 
pepper. 



26. Xanthomonas nakatae (Okabe) 
Dowson. {Bacterium nakatae Type B, 
Okabe, Jour. Soc. Trop. Agr., Formosa, 
5, 1933, 161 ; Phylomonas nakatae Burk- 
holder, in Manual, 5th ed., 1939, 154; 
Dowson, Trans. Brit. Mycol. Soc, 26, 
1943, 12.) Named for Nakata, the Japa- 
nese plant pathologist. 

Rods: 0.3 to 0.4 by 1.1 to 2.5 microns. 
Capsules. Motile with a polar flagellum. 
Gram-negative. 

Gelatin: Liquefaction. Brown color. 

Beef -extract agar colonies: Amber yel- 
low, round, smooth, glistening, margins 
entire. Brown. 

Broth : Moderate turbidity with yellow 
ring. Medium turns brown. 

Milk: Casein is precipitated and di- 
gested. Tyrosine crystals. Brown color. 

Nitrites not produced from nitrates. 

Indole not formed. 

Slight amount H2S produced. 

Acid but not gas from glucose, sucrose, 
maltose and lactose. 

Starch: Strong diastatic action. 

Optimum temperature 30° to 32°C. 
Maximum 39°C. Minimum 10°C. 

No growth in beef extract broth plus 
2 per cent salt. 

Aerobic. 

Distinctive character: Differs from 
Type A in that it produces a brown pig- 
ment in culture. (Description of Type A 
not seen.) 

Source : Isolated from water-soaked to 
brown leaf spots on jute. 

Habitat : Pathogenic on jute, Corchoriis 
capszdaris. 

27. Xanthomonas papavericola (Bryan 
and McWhorter) Dowson. {Bacterium 
papavericola Bryan and McWhorter, 
Jour. Agr. Res., 40, 1930, 9; Phytomonas 
papavericola Bergey et al., Manual, 4th 
ed., 1934, 266; Dowson, Cent. f. Bakt., 
IIAbt.,iOO, 1939,190.) FromL. papaver, 
poppy; -cola, dweller; M. L. Papaver, a 
generic name. 

Rods: 0.6 to 0.7 by 1 to 1.7 microns. 
Chains. Capsules. Motile with a single 
polar flagellum. Gram-negative. 



FAMILY PSEUDOMONADACEAE 



165 



Gelatin : Liquefaction. 

Beef agar colonies : Mustard yellow to 
primuline yellow, circular, margins 
entire. 

Broth : Turbidity prompt with a yellow 
ring and an incomplete pellicle. 

Milk : Soft coagulation, peptonization 
and production of tyrosine crystals. 

Nitrates : A weak reaction for nitrites 
after 10 days. 

Indole not formed. 

Hydrogen sulfide is produced. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 5:?, 1942, 600). 

Acid but not gas from glucose, galac- 
tose, fructose, sucrose, lactose, maltose, 
glycerol and mannitol. 

Starch is hydrolyzed. 

Optimum temperature 25° to W°C. 
Maximum 35°C. 

No growth in broth plus 5 per cent salt. 

Aerobic. 

Source: Isolated from black spots on 
leaves, buds and pods of poppy. 

Habitat : Pathogenic on poppy, Papaver 
rhoeas. 

28. Xanthomonas alfalfae (Riker 
et al.) Dowson. {Bacterium alfalfae 
Riker, .Tones and Davis, Jour. Agr. Res., 
51, 1935, 177; Phylomonas alfalfae Riker 
ot al., ihid.; Pseudomonas alfalfae Riker 
ot al., ihid.; Dowson, Trans. Brit. Alycol. 
Soc, 26, 1943, 11.) From Spanish, of 
alfalfa. 

Rods: 0.45 by 2.4 microns. Motile 
with a polar fiagellum. Gram-negative. 

Gelatin: Liquefied. 

Nutrient agar stroke : Growth abun- 
dant, filiform, smooth, glistening, butyr- 
ous, pale yellow. 

Broth: Turbid in 24 hours. Light 
sediment. 

Milk: Casein is precipitated and di- 
gested. 

Ammonia formed slowly in a nitrate 
medium. 

Carbohydrates : No acid in yeast broth 
plus sugars. 

Starch is hydrolyzed. 

Aerobic. 



Optimum temperature 24° to 32°C. 
Maximum below 36°C. Minimum below 
4°C. 

Source : Six single cell cultures isolated 
from diseased alfalfa. 

Habitat : Pathogenic on the leaves of 
alfalfa, Medicago sativa. 

29. Xanthomonas acernea (Ogawa) 
comb. nov. {Pseudomonas acernea Ogawa, 
Ann. Phyt. Soc. Japan, 7, 1937, 123; 
Phylomonas acernea Ark, Phytopath., 
29, 1939, 968.) From L. acerneus, of 
the maple. 

Rods: 0.2 to 0.6 by 0.5 to 1.2 microns. 
Motile with one polar fiagellum. Gram- 
negative. 

Gelatin: Liquified. 

Agar colonies : Round, smooth, convex, 
white to citron yellow, glistening, trans- 
lucent with amorphous structure. 

Broth : Turbid. 

Milk: Slowly cleared, slightly acid. 
No coagulation. 

Nitrites produced from nitrates. 

Hydrogen sulfide produced. 

No gas produced in peptone water plus 
sugars. 

Starch not hydrolyzed. 

Optimum temperature about 32°C. 
Thermal death point 59°C. 

Aerobic. 

Source : From diseased leaves of Acer 
trifidum in Japan. 

Habitat : Causes a disease in Acer spp. 
and in Aesculus turbinaia and Koelren- 
teria panicvlata. 

30. Xanthomonas carotae (Kendrick) 
Dowson. {Phylomonas carotae Ken- 
drick, Jour. Agr. Res., 49, 1934, 504; 
Pseudomonas carotae Kendrick, ibid. ; 
Dowson, Cent. f. Bakt., II Abt., 100, 
1939, 190.) From L. carota, the carrot. 

Rods: 0.42 to 0.85 by 1.38 to 2.75 
microns. Motile with 1 or 2 polar flagella. 
Gram-negative. 

Gelatin: Liquefied. 

Potato glucose agar: Colonies round, 
smooth, glistening, margins entire, straw 
yellow in color. 



166 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Milk : Casein precipitated and milk 
cleared; alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

Acid, no gas, from glucose, d-galac- 
tose, xylose, rf-mannose, Z-arabinose, 
sucrose, lactose, raffinose, trehalose, 
d-mannitol and glycerol. No acid from 
maltose and rhamnose. 

Starch not hydrolyzed. 

Optimum temperature 25° to 30°C. 

Tolerates 4 per cent salt at pH 7. 

Aerobic. 

Source : Two original isolations from 
diseased carrots and a reisolation from 
inoculated carrots were used for the 
description. 

Habitat: Pathogenic on leaves of 
Daucus caroin var. saliva. 

31. Xanthomonas hederae (Arnaud) 
Dowson. {Bacterium hederae Arnaud, 
Compt. rend. Acad. Sci., Paris, 171, 
1920, 121; Phytomonas hederae Burk- 
holder and Guterman, Phj^topath., 22, 
1932, 783; Dowson, Cent. f. Bakt., II 
Abt., 100, 1939, 190.) From L. hedera, 
ivy; M. L. Hedera, a generic name. 

Description taken from Burkholder 
and Guterman {loc. cit.). 

Rods: 0.6 by 2.13 microns. Motile 
with a single polar flagellum. Gram- 
negative. 

Gelatin: Liquefied. 

Beef-extract-agar slants: Growth good, 
filiform, amber yellow, butyrous. 

Broth: Turbid. 

Milk: Casein is precipitated and di- 
gested. Milk becomes alkaline. 

Nitrites not produced from nitrates. 

Hydrogen sulfide is formed. 

Indole not formed. 

Not lipolytic (Starr and Burkholder, 
Phytopath., 32, 1942, 600). 

Acid from glucose, fructose, galactose, 
xylose, sucrose, lactose and glycerol. 
Alkali from salts of acetic, citric, lactic, 
malic and succinic; acids. The following 
are not utilized : arabinose, rhamnose, 
maltose, salicin, starch, cellulose and 
formic acid. 



Aerobic, facultative. 

Source : Isolated from diseased ivy 
leaves. 

Habitat : Pathogenic on ivy, Hedera 
helix. 

32. Xanthomonas phormicola (Taki- 
moto) Dowson. {Bacterium phormicola 
Takimoto, Jour. Plant Protect., 20, 1933, 
777; Phytomonas -phormicola Burkholder, 
in Manual, 5th ed., 1939, 159; Dowson, 
Trans. Brit. Mycol. Soc, 26, 1943, 12.) 
From M. L. Phormium, a generic name. 

Description translated by Dr. K. 
Togashi . 

Rods: 0.5 to 0.6 by 1 to 2 microns. 
Motile, with a single flagellum. Gram- 
negative. 

Gelatin : Liquefied. 

Agar colonies : Light yellow, then waxy 
yellow; butyrous, then viscid. 

Broth : Turbid, pellicle formed. 

Milk: Casein coagulated slowly and 
precipitated, then digested. Alkaline. 

Nitrites not produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced. 

No gas from sucrose, glucose, lactose 
and glycerol. 

No acid from various sugars in broth. 

Optimum temperature about 29°C. 
Maximum 39°C. Minimum about 0°C. 

Aerobic. 

Source : Species isolated from New 
Zealand flax, Phormium tenax. 

Habitat: Causes a leaf stripe of Phor- 
mium tenax. 

33. Xanthomonas geranii (Burkholder) 
Dowson. {Phytomonas geranii Burk- 
holder, Phytopath., 27, 1937, 560; Dow- 
son, Cent. f. Bakt., II Abt., 100, 1939, 
190.) From Greek, geranos, crane; M. 
L. Geranium , a generic name. 

Rods: 0.75 to 2.0 microns. Motile 
with a single polar flagellum. Gram- 
negative. 

Gelatin: Liquefied. 

Beef -extract agar slants : Moderate to 
good filiform growth, glistening, primu- 
line yellow. Develops in 24 hours. 



FAMILY PSEtJDO:MONADACEAE 



167 



Broth : Turbid in 24 hours. No pellicle 
but a moderate sediment. 

Milk : Becomes clear with a heavy 
casein precipitate. Peptonization with 
crystal formation. 

Nitrates reduced to ammonia. 

Indole not formed. 

Hydrogen sulfide formed. 

Lipolytic (Starr and Burkholder, Phy- 
topath., 32, 1942, 600). 

Acid from glucose, galactose, fructose, 
xylose, rhamnose, lactose, sucrose, raf- 
finose and glycerol. Alkaline reaction 
from salts of citric, malic, malonic and 
succinic acid. No growth in arabinose 
or formic, hippuric, maleic or tartaric 
acid. 

Starch not hydrolyzed. 

Aerobe. 

Distinctive characters : Pathogenic on 
Geranium spp., not on the house gera- 
nium, Pelargonium hortorum. In culture 
similar to Xanlhomonas pelargonii. 

Source : Three cultures isolated from 
Geranium- sangnineurn. 

Habitat : Pathogenic on Geranium san- 
guineum, G. maculatum, G. pratense and 
G. sylvaticum. 

34. Xanthomonas antirrhini (Taki- 
moto) Dowson. {Psendomonas anti- 
rrhini Takimoto, Bot. Mag. Tokyo, 34, 
1920, 257; Bacterium antirrhini Elliott, 
Man. Bact. Plant Path., 1930, 93; Phyto- 
monas antirrhini Magrou, in Hauduroy 
et al., Diet. d. bact. path., Paris, 1937, 
331; Dowson, Trans. Brit. Mycol. Soc, 
26, 1943, 11.) From Gr. antirrhinum, 
snapdragon; M. L. Antirrhinum, a 
generic name. 

Description from Elliott (loc.cit.). 

Rods: 0.3 to 0.4 by 0.8 to 1.2 microns. 
Motile with polar flagella. Capsules. 
Gram-negative. 

Gelatin: Liquefied. 

Agar colonies: Round, glistening, 
white, later yellow. 

Milk: Coagulated and casein digested. 

Nitrites are produced from nitrates. 

No gas produced. 

Aerobic. 



Optimum temperature 26° to 27°C. 
Maximum 34°C. 

Habitat: Causes a leaf spot of Anti- 
rrhinum majus. 

35. Xanthomonas heterocea (Vzoroff) 
comb. nov. {Phytomonas heterocea 
Vzoroff, Bull. North Caucasian Plant 
Prot. Sta. Roztoff-on-Don, 6-7, 1930, 
263; Bacterium heteroceum Burgwitz, 
Phytopathogenic bacteria, Leningrad, 
1935, 135.) From Gr. hcterus, another, 
different. 

Description taken from Rev. App. 
Myc, 10, 1931, 628. 

Rods : 0.4 to 0.6 by 1.0 to 2.0 microns. 
Motile. Gram-negative. 

Gelatin : Slow liquefaction. 

Agar colonies : Round, convex, smooth, 
semi-transparent, glistening, yellow to 
amber, 2 mm. in diameter. Pitted sur- 
face . 

Milk: No coagulation. At first acid, 
later alkaline. 

Nitrites produced from nitrates. 

Indole not formed. 

Hydrogen sulfide produced. 

Acid from glucose, galactose, arabi- 
nose, xylose, sucrose, maltose, salicin, 
glycerol and mannitol . Does not ferment 
lactose, inulin, ethyl alcohol, esculin, 
adonitol or dulcitol. 

Optimum temperature 25° to 30°C. 

Source : Isolated from diseased tobacco 
in the North Caucasus. 

Habitat : Pathogenic on Nicotiana 
tabacum. 

36. Xanthomonas gummisudans (Mc- 
Culloch) Starr and Burkholder. (Bac- 
terium gummisiulans McCulloch, Phyto- 
path., H, 1924, 63; also Jour. Agr. Res., 
27, 1924, 229; Pseudomonas gummisudans 
Stapp, in Sorauer, Handb. d. Pflanzen- 
krank., 2, 5 Aufl., 1928, 54; Phytomonas 
gummisudans Bergey et al., Manual, 2nd 
ed., 1925, 201; Starr and Burkholder, 
Phytopath., 32, 1942, 600.) From L. 
gummi, gum; sudans, sweating, dripping. 

Rods : 0.6 to 0.8 by 1 to 2.8 microns. 



168 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Capsules. Motile with a polar flagellum. 
Gram-negative. 

Gelatin : Liquefied. 

Beef -peptone agar colonies : Amber 
yellow, circular, transparent, smooth, 
with definite margins. 

Broth : Moderately turbid with a yel- 
low ring. 

Milk : Soft curd which is digested with 
formation of tyrosine crystals. 

Nitrites not produced from nitrates. 

Indole not produced. 

Hydrogen sulfide produced. 

Lipolytic (Starr and Burkholder, loc. 
cit.). 

Acid from glucose and sucrose. 

Optimum temperature 30°C. Maxi- 
mum Se'C. Minimum 2°C. 

Aerobic. 

Source : From gummy lesions on gladio- 
lus leaves. 

Habitat : Pathogenic on leaves of glad- 
ioli. 

37. Xanthomonas lactucae (Yama- 
moto) Dowson. (Bacterium lactucae 
Yamamoto, Jour. Plant Protect., 21, 
1934, 532; Phytomonas lactucae Bergey 
et al.. Manual, 5th ed., 1939, 163; Dow- 
son, Trans. Brit. Mycol. Soc, 26, 1943, 
12.) From L. lactuca, lettuce; M. L. 
Lactuca, a generic name. 

Description translated by Dr. K. 
Togashi . 

Rods : 0.6 to 0.8 by 1.75 to 2.8 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin : Liquefaction slow. 

Agar colonies: Circular, convex, mar- 
gin entire, surface smooth, wet-shining, 
yellow. 

Broth: Turbid. Ring and pellicle. 

Milk: Slow peptonization. 

Nitrites not produced from nitrates. 

Indole not produced. 

Hydrogen sulfide produced. 

Acid, no gas, from glucose, sucrose, 
and lactose in bouillon; no acid from 
glycerol in bouillon. 

Optimum temperature 28°C. Maxi- 
mum Z5°C. Minimum below 2''C. 



Aerobic. 

Source : Isolated from leaf spot of 
lettuce. 

Habitat : Pathogenic on leaves of as- 
paragus lettuce, Lactuca saliva var. an- 
gustata. 

38. Xanthomonas nigromaculans 
(Takimoto) Dowson. (Bacterium nigra - 
m.aculans Takimoto, Jour. Plant Protect., 
Tokyo, H, 1927, 522; Phytomonas nigro- 
maculans Magrou, in Hauduroy et al., 
Diet. d. Bact. Path., Paris, 1937, 387; 
Dowson, Trans. Brit. Mycol. Soc, 26, 
1943, 12.) From L. niger, black; macu- 
lans, spotting. 

Description translated by Dr. K. 
Togashi . 

Rods: 0.6 to 0.9 by 1.5 to 2.8 microns. 
Motile with 1 or 2 polar flagella. Gram- 
negative. 

Gelatin: Liquefaction. 

Agar colonies : Yellow, circular, margins 
entire, smooth, glistening. 

Broth : Growth moderate with yellow 
pellicle. 

Milk : Coagulation and digestion of the 
casein. 

Nitrites not produced from nitrates. 

Indole not produced. 

No acid or gas from glucose, sucrose, 
lactose, mannitol and glycerol in peptone 
water. 

Optimum temperature 27° to 28°C. 
Maximum 33°C. Minimum O^C. 

Aerobic. 

Source : Isolated from lesions on leaf 
and petioles of burdock. 

Habitat: Pathogenic on leaves and 
petioles of Arctium lappa, the burdock. 

39. Xanthomonas oryzae (Uyeda and 
Ishiyama) Dowson. {P seudomonas 
oryzae Uyeda and Ishiyama, Proc. Third 
Pan-Pacific Sci. Congr., Tokyo, 2, 1926, 
2112; Bacterium- oryzae Nakata, see 
Elliott, Man. Bact. Plant Path., 1930, 
172; Phytomonas oryzae Magrou, in 
Hauduroy et al.. Diet. d. Bact. Path., 
Paris, 1937, 388; Dowson, Trans. Brit. 



FAMILY PSEUDOMONADACEAE 



169 



Mycol. Soc, 26, 1943, 12.) From Gr. 
oryza, rice; M. L. Oryza, a generic name. 

Probable synonym : Pseudomonas ito- 
ana Tochinai, Ann. Phytopath. Soc. 
.Japan, 2, 19.32, 456; Bacterium itoanum 
Burgwitz, Phytopathogenic Bacteria, 
Leningrad, 1935, 74; Phytomonas itoana 
Magrou, in Hauduroy et al.. Diet. d. 
Bact. Path., Paris, 1937, 370. 

Rods: 0.5 to 0.8 by 1.0 to 2.0 microns. 
Motile with a polar flagellum. Gram- 
negative. 

Gelatin: No liquefaction. 

Nutrient agar colonies : Round, smooth, 
glistening, wax yellow. 

Milk: Slightly acid. 

Nitrites are not produced from nitrates. 

Hydrogen sulfide produced. 

Acid but no gas from glucose, lactose 
and sucrose. 

Optimum temperature 26° to 30°C. 

Strict aerobe. 

Source : Isolated from a leaf blight of 
rice. 

Habitat : Pathogenic on rice, Oryza 
saliva. 

40. Xanthomonas celebensis (Gau- 
mann) Dowson. {Pseudomonas celeben- 
sis Gaumann, Ztschr. f. Pflanzenkrank., 
S3, 1923, 11; Meded. Inst, voor Planten- 
ziek., Buitenzorg, 59, 1923, 17; Bacterium 
celebense Elliott, Man. Bact. Plant Path., 
1930, 108; Phytomonas celebensis Magrou, 
in Hauduroy et al., Diet. d. Bact. Path., 
Paris, 1937, 343; Dowson, Trans. Brit. 
Mycol. Soc, 26, 1943, 11.) From M. L. 
of the island Celebes. 

Rods : 0.9 by 1.5 microns. Motile by a 
polar flagellum. Gram -negative. 

Agar colonies: Grayish yellow. 

Broth: Thin pellicle. 

Milk: Coagulated and cleared. 

Nitrites not produced from nitrates. 

Sodium selenite: Brick red. 

Starch is hydrolyzed. 

Source : From vascular bundles of 
diseased bananas in Celebes. 

Habitat : Causes the blood disease of 
banana. 



41. Xanthomonas panici (Elliott) 
comb. nov. {Bacterium panici Elliott, 
Jour. Agr. Res., 26, 1923, 157; Pseudo- 
monas panici Stapp, in Sorauer, Handb. 
d. Pflanzenkrank., 2, 5 Aufl., 1928, 27; 
Phytomonas panici Bergey et al., Man- 
ual, 3rd ed., 1930, 269.) From M. L. 
Panicum, a generic name. 

Rods : 0.69 by 1.66 microns. Capsules. 
Motile by 1 or rarely 2 polar flagella. 
Gram-negative. 

Gelatin : Liquefaction slow. 

Beef agar colonies: Round, white, 
smooth, glistening, margins at first entire, 
later undulate. 

Broth : Moderate turbidity in 24 hours. 
Thin pellicle. Medium brownish. 

Milk: Alkaline and clears. 

Nitrites are produced from nitrates. 

Indole not produced. 

Hydrogen sulfide produced. 

No gas from carbohydrates. 

Starch: Hydrolysis moderate. 

Optimum temperature 33°C. Maxi- 
mum 45°C. Minimum 5°C. 

Optimum pH 6.15 to 6.3. pH range 
5.4 to 10.0. 

Aerobic. 

Distinctive characters : Differs from 
Pseudomonas andropogoni in that it 
liquefies gelatin, produces nitrites from 
nitrates, and does not infect sorghum and 
broom corn. 

Source : Isolation from water soaked 
lesions on leaves, sheaths and culms of 
millet collected in Wisconsin and in S. 
Dakota. 

Habitat : Pathogenic on proso millet, 
Panicum m.iliaceum, 

42. Xanthomonas proteamaculans 
(Paine and Stansfield) comb. nov. {Pseu- 
domonas proteamacidans Paine and Stans- 
field, Ann. Appl. Biol., 6, 1919, 38; 
Phytomonas proteamacidans Bergey et 
al.. Manual, 3rd ed., 1930, 247; Bacterium 
proteamacidans Elliott, Man. Bact. Plant 
Path., 1930, 186.) From M. L. Protea, 
a generic name; maculans, spotting. 

Rods : 0.6 to 0.8 by 0.8 to 1.6 microns. 



170 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Motile with 1 to 3 polar flagella. Gram- 
positive. 

Gelatin : Liquefaction. 

Agar slant : Growth wet-shining, dirty 
white with a faint yellow tinge. 

Broth: Turbid in 24 hours. Slight 
ring. 

Milk : Acid with soft curd after 2 days. 
Later a separation of whey. 

Nitrites are produced from nitrates. 

Acid and gas from glucose, sucrose and 
mannitol. No acid or gas from lactose. 

Starch: Slight hydrolysis. 

Source : Repeated isolation from a leaf- 
spot of Proica in England. 

Habitat : Pathogenic on Protea cy- 
nar aides. 

43. Xanthomonas manihotis (Arthaud- 
Berthet) comb. nov. {Bacillus manihotus 
Arthaud-Berthet by Bondar, Chacaras 
and Quintaes 5(4), 1912, 15; Bacillus 
manihot Bondar (and Arthaud-Berthet), 
Bol. Agric, Sao Paulo, 16, 1915, 513; 
Bacterium m.anihoius Drummond and 
Hipolito, Ceres, 2, 1941, 298; Phytomonas 
manihotis Viegas, Rev. d. Agr., Pieraci- 
caba, 15, 1940, 475.) From M. L. Mani- 
hotus, a generic name. 

Description from Burkholder, Phyto- 
path., 32, 1942, 147. 

Rods : 0.35 to 0.93 by 1.4 to 2.8 microns. 
Gram-negative and mostly non-motile. 
One isolate showed a few cells with 1 
polar flagellum. Amaral (Instit. Biol., 
Sao Paulo, Arq., IS, 1942, 120) states that 
the species is motile with one polar fla- 
gellum. 

Gelatin: Liquefaction. 

Beef -extract-peptone agar : Streaks 
raised, ivory-color, smooth, shiny, with 
edges entire. 

Potato-glucose agar : Growth abun- 
dant, white to hyaline, very mucoid. 

Broth : Turbid with a whitish granular 
ring. 

Litmus milk : Litmus reduced and 
milk clears. With return of color, litmus 
is purple. 

Indole not formed. 

Hydrogen sulfide is formed. 



Nitrites produced from nitrates 
(Drummond and Hipolito, loc. cit.). 

Asparagine not used as a nitrogen and 
carbon source. No growth in nitrate 
synthetic broth. 

Weak growth but slight acid production 
in synthetic medium plus glucose, 
d-galactose, d-fructose, d-xylose, maltose 
and sucrose. No growth in rhamnose, 
1-arabinose, d-lactose, glycerol, mannitol 
and salicin. Good growth with alkaline 
reaction in same medium plus salts of the 
following acids: acetic, citric, malic, 
maleic and succinic. The salts of formic, 
hippuric, lactic and tartaric acids were 
not utilized. 

Starch not hydrolyzed. Amaral {loc. 
cit.) finds hydrolysis. 

Lipolytic action slight. 

Aerobic. 

Optimum temperature 30°C. Maxi- 
mum 38°C. Minimum 5°C. 

Source : First isolated from the cassava, 
Manihotus utilissima in Brazil. 

Habitat : Produces a wilt disease on 
various species of Manihotus. 

44. Xanthomonas rubrisubalbicans 

(Christopher and Edgerton) comb. nov. 
{Phytomonas rubrisubalbicans Christo- 
pher and Edgerton, Jour. Agr. Res., 4^, 
1930, 266; Bacterium rubris2tbalbicans 
Burgwitz, Phytopathogenic Bacteria, 
Leningrad, 1935, 105.) From L. ruber, 
red; subalbicans, nearly white. 

Rods: Short with polar flagella. Cap- 
sules. Gram-negative. 

Gelatin: No liquefaction. 

Bacto-glucose agar colonies: Circular, 
glistening, viscid, milky gray to buff. 
Margins translucent, entire. 

Broth : Turbid after 24 hours. Pellicle 
and a ropy sediment. 

Indole produced. 

Hydrogen sulfide produced. 

No gas from carbohydrates. 

Starch hydrolyzed. 

Optimum temperature 30°C. 

Optimum pH 6.8 to 8.0. 

Source : Isolated many times from mot- 
tled stripe of sugar cane in Louisiana. 



FAMILY PSEUDOMOXADACEAE 



171 



Habitat : Pathogenic on sugar cane, 
Johnson's grass and sorghum. 

45. Xanthomonas cannae (Bryan) 
comb. nov. {Bacterium cannae Bryan, 
Jour. Agr. Res.,^i, 1921, \b2;Phytomonas 
cannae Bergey et al., Manual, 1st ed., 
1923, 188; Pseudomonas cannae Stapp, in 
Sorauer, Handb. d. Pflanzenkrank., 2, 
5 Aufl., 1928, 65.) From Gr. canna, a 
reed; M. L. Canna, a generic name. 

Rods: 0.5 to 0.7 by 1.0 to 2.0 microns. 
Motile with 1 to 3 polar flagella. Cap- 
sules. Gram -negative. 

Gelatin: Slow liquefaction. 

Agar streaks: Filiform, white, moist, 
with thin margins and granular centers. 

Broth: Turbid, heavy sediment. 

Milk: Alkaline and clears. 

Nitrites are produced from nitrates. 

Indole not produced. 

Hydrogen sulfide produced. 

Optimum temperature 35°C. Maxi- 
mum 40°C. Minimum 5°C. 

Aerobic. 

Source : Isolated from diseased canna 
leaves collected in Washington, D. C. 
and in Illinois. 

Habitat : Causes disease in Canna 
indica. 

46. Xanthomonas zingiberi (LVeda) 
comb. nuv. (Uyeda, Cent. f. Bakt., II 
Abt., 17, 1907, 383; Pseudomonas zingi- 
beri Uyeda, Rept. Imp. Agr. Exp. Sta., 
Japan, No. 35, 1908, 114; Bacterium 
zingiberi Nakata, see Elliott, Man. Bact. 
Plant Path., 1930, 266; Phytomonas zingi- 
beri Magrou, in Hauduroy, et al., Diet. d. 
Bact. Path., Paris, 1937, 437.) From L. 
zingiberis, ginger; M. L. Zingiber, a 
generic name. 

Description from Stapp, in Sorauer, 



Handb. d. Pflanzenkrank., 2, 5 Aufl., 
1928, 65. 

Rods : 0.5 to 1.1 by 0.75 to 1.8 microns. 
Non-motile at first, later a polar flagel- 
lum. Gram-negative. 

Gelatin: Liquefaction. 

Agar colonies : White. 

Milk: Coagulation and peptonization of 
the casein. 

Nitrites are produced from nitrates. 

Indole not formed. 

Hydrogen sulfide is formed. 

No gas from glucose. 

Optimum temperature 28°C. Maxi- 
mum 40°C. Minimum 5°C. 

Source : Isolated from ginger plant 
showing a rot at the base of the sprouts. 

Habitat: Pathogenic on ginger, Zingi- 
ber officinale. 

47. Xanthomonas conjaci (Uyeda) 
comb. nov. (Pseudomonas conjac Uyeda, 
Bot. Mag. Tokyo, 24, 1910, 182; Bac- 
terium conjac Elliott, Man. Bact. Plant 
Path., 1930, 121; Phytomonas conjac 
Magrou, in Hauduroy et al., Diet. d. 
Bact. Path., Paris, 1937, 347.) From 
M. L. conjac, the specific name of the 
plant which this species attacks. 

Description from Elliott (loc.cit.). 

Rods: 0.75 to 1.0 by 1.5 microns. 
Motile with 1 to 4 polar flagella. Gram- 
positive. 

Gelatin colonies : Circular to irregular, 
light yellow. 

Broth: Pellicle formed. 

Milk: Coagulation. 

Conjac: Liquefied. 

Nitrites produced from nitrates. 

Indole produced. 

Hydrogen sulfide produced. 

Gas from glucose. 

Favorable temperature 24°C. 

Habitat : Pathogenic on Amorphophal- 
lus konjac. 



Appendix I:* The following organisms placed in the genus Psei^domottas apparently 
belong in Xanthomonas. Some may even be plant pathogens although they were 

* Prepared by Prof. Robert S. Breed, New York State Experiment Station, Geneva, 
New York, July, 1943. 



172 MANUAL OF DETERMINATIVE BACTERIOLOGY 

isolated from water, soil and similar sources. Pigment is usually yellow and is not 

water-soluble. 

Key to yellow and other chromogenic species in genus Pseudomonas. 

1. Colonies yellow. 

a. Gelatin liquefied. 

b. Nitrites produced from nitrates. 

c. Acid and gas produced from glucose. 

1 . Pseudomonas fermentans . 
cc. Acid but no gas from glucose. 

2. Pseudomonas trifolii. 

3. Pseudomonas xanthe. 
ccc. Action on glucose not recorded. 

4. Pseudomonas caudata. 
bb. Nitrites not produced from nitrates. 

c. Litmus milk acid or ferment lactose. 

5. Pseudomonas perlurida. 

6. Pseudomonas iridescens. 
cc. Litmus milk not coagulated. Yellow sediment. 

7. Pseudomonas turcosa. 
ccc. Litmus milk slimy, alkaline. 

8. Psezidomonas ochracea. 
aa. No liquefaction of gelatin. 

b. Nitrites produced from nitrates. 

c. Litmus milk, slow coagulation. 

9. Pseudomonas cerevisiae. 
cc. Litmus milk, acid but no digestion. 

10. Pseudomonas arguta. 
ccc. No growth in litmus milk. 

11. Pseudomonas subcreta. 
cccc. Action on litmus milk, not recorded. 

12. Pseudomonas pictorjun. 
bb. Nitrites not produced from nitrates. 

c. Butter colored pellicle on litmus milk. 

13. Pseudomonas lacunogenes. 
cc. No surface pellicle. 

14. Pseudomonas segnis. 

2. Colonies on gelatin blue center surrounded by yellow zone with peripheral 

green zone, 
a. Gelatin liquefied. 

b. Nitrites produced from nitrates. 

15. Pseudomonas lemonnieri. 

1. Pseudomonas fermentans von Wol- with rounded ends, occurring singly and 

zogen Kiihr. (von Wolzogen Kiihr, in pairs. Motile, with a single or occa- 

Cent. f. Bakt., II Abt., 85, 1932, 228; sionally 2 or 3 polar flagella. Gram- 

Flavobacterium fermentans Bergey et al., negative. 

Manual, 4th ed., 1934, 155.) From Gelatin colonies: Circular, grayish, 

Latin, fermento, to ferment. with rapid liquefaction. 

Rods: 0.4 to 0.6 by 1.7 to 3.4 microns. Gelatin stab : Liquefaction crateriform. 



FAMILY PSEUDOMONADACEAE 



173 



Agar colonies : Circular, slightly con- 
vex, opaque, gray by reflected, and 
light-brown by transmitted light. 

Agar slant : Gray, becoming yellowish. 

Broth: Turbid with pellicle. 

Litmus milk: Acid. 

Potato : Gray to yellowish growth. 

Indole is formed. 

Nitrites produced from nitrates. 

Acid and visible gas from glucose, 
lactose and sucrose. 

Acetylmethylcarbinol is formed. 

Ammonia is formed from peptone and 
asparagin. 

Hydrogen sulfide is formed. 

Starch is hydrolyzed. 

Lipase is formed. Catalase positive. 

Aerobic, facultative. 

Optimum temperature 37°C. 

Distinctive character; Produces gas in 
lactose fermentation tubes. 

Source : Ten cultures from the larvae 
of a midge (Chironomtis plumosus) and 
from filtered water. 

Habitjit: Unknown. 

2. Pseudomonas trifolii Huss. (Huss, 
Cent. f. Bakt., II Abt., 19, 1907, 68; 
Flavobacterium trifolii Bergey et al., 
Manual, 1st od., 1923, 111.) From Latin, 
tres (Iri-), three; Jolivm, leaf; M. L. 
Trifolium, clover. 

Possible synonym : Bacillus annulatus 
Wright. (Wright, Memoirs Nat. Acad. 
Sci., 7, 1895, 443; Pseudomonas annidata 
Chester, Man. Determ. Bact., 1901, 315; 
Relationship to Bacillus annulatus Zim- 
mermann uncertain. Die Bakt. unserer 
Trink- und Nutzwiisser, Chemnitz, II 
Reihc, 1890, 30; Flavobacterium annula- 
lum Bergey et al., Manual, 1st ed., 1923, 
110.) 

According to Mack (Cent. f. Bakt., II 
Abt., 95, 1936, 218) the following organism 
is to be regarded as identical with Pseudo- 
monas trifolii: Bacillus mesentericus 
aureus Winkler (Cent. f. Bakt., II Abt., 
5, 1899, 577) regarded by Burri (Cent. f. 
Bakt., II Abt., 10, 1902, 756) and Dtiggeli 
(Cent. f. Bakt., II Abt., 12, 1904, 602) 



as identical with the organism which 
Duggeli (loc. cit.) names Bacterium 
herbicola aureum. The organism studied 
as Bacterium herbicola by Hiittig (Cent, 
f. Bakt., II Abt., 84, 1931, 231) is not 
regarded as identical with the Burri and 
Diiggeli organism by Mack. Beijerinck 
(Cent. f. Bakt., II Abt., IB, 1905, 366) 
states that Bacillus herbicola of Burri 
and Diiggeli is identical with his Bacillus 
anglomerans (Botan. Ztg., 1888, 749). 
If so, this binomial has priority. 

Rods : 0.5 to 0.7 by 0.75 to 2.0 microns, 
occurring singly, in pairs and in chains. 
Motile, possessing a single polar flagellum. 
Gram-negative. 

Gelatin colonies : Convex, smooth, 
moist, glistening, grayish -yellow. 

Gelatin stab: Napiform liquefaction. 

Agar colonies: Small, circular, grayish, 
becoming brownish-yellow. 

Agar slant : Yellowish, becoming brown- 
ish-yellow streak, lacerate margin. 

Broth: Turbid, with grayish-yellow 
pellicle and sediment. 

Litmus milk: Slowly coagulated; alka- 
line; with yellow ring. 

Potato: Thick, yellowish, fiat, smooth, 
glistening. 

Hj'drogen sulfide produced. 

Indole is formed. 

-\cid from glucose, sucrose, xylose, 
arabinose, and mannitol. No acid from 
lactose. 

Nitrites produced from nitrates. 

Cultures have an agreeable odor. 

Volutin formed. 

Aerobic, facultative. 

Optimum temperature 33° to 35°C. 

Source: Isolated from clover hay. 

Habitat : Evidently a common organism 
on the leaves of plants. 

3. Pseudomonas xanthe Zettnow. 
(Zettnow, Cent. f. Bakt., I Abt., Orig., 
77, 1915, 220; Flavobacterium zettnowii 
Bergey et al.. Manual, 1st ed., 1923, 
112; Flavobacterium xanthium (sic) Ber- 
gey et al.. Manual, 3rd ed., 1930. 145.) 
From Gr. xanthus, yellow. 



174 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Rods: 0.5 to 0.6 by 0.4 to 1.4 microns. 
Motile, possessing a single or occasionally 
two or more very long (20 microns) polar 
flagella. Gram-negative. 

Gelatin colonies: Circular, yellow, 
granular. 

Gelatin stab : Pale-yellow surface 
growth. Brownish yellow under surface 
colonies. Saccate liquefaction. 

Agar slant : Dark yellow, glistening, 
with dark yellow sediment in water of 
condensation. Pigment not water-sol- 
uble. 

Broth: Turbid. 

Litmus milk: Slightly acid. Litmus 
reduced. 

Potato : Grayish yellow to brownish 
growth. 

Indole formed. 

Nitrites are produced from nitrates. 

Acid formed in glucose. 

Starch hydrolyzed. 

Blood serum not liquefied. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Source: Air contamination. 

4. Pseudomonas caudata (Wright) 
Conn. {Bacillus caudatus Wright, 
Memoirs Nat. Acad. Sci., 7, 1895, 444; 
Bacterium caudatus Chester, Annual 
Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 
107; Conn, Jour. Agr. Res., 16, 1919, 313; 
Flavobacterium caudatum Bergey et al., 
Manual, 1st ed., 1923, 109.) From 
Latin, cauda, tail. 

Rods : Long, granular, slender, occurring 
singly, in pairs and in chains. Appear 
like cocci in old cultures. Motile, pos- 
sessing a polar flagellum (Conn). Gram- 
negative. 

Gelatin colonies: Yellow, translucent, 
smooth, undulate. 

Gelatin stab: Villous growth in stab. 
Crateriform liquefaction. 

Agar slant: Yellow to orange, glisten- 
ing, translucent, slightly spreading. 
May lose power to form pigment. 

Broth: Turbid, with yellow sediment. 

Litmus milk: Unchanged. 

Potato: Dark yellow, raised, rough, 
spreading. 



Indole not formed. 

Nitrites and ammonia produced from 
nitrates. 
Ammonia produced from peptone. 
Starch is digested. 
Aerobic, facultative. 
Optimum temperature 25°C. 
Habitat : Water. 

5. Pseudomonas perlurida Kellerman 
et al. (Kellerman, McBeth, Scales and 
Smith, Cent. f. Bakt., II Abt., 39, 1913, 
516; also McBeth, Soil Sci., 1, 1916, 472; 
Ccllulomonas perlurida Bergey et al., 
Manual, 1st ed., 1923, 163.) 

Rods: 0.4 by 1.0 micron. Motile with 
one to three polar flagella. Gram- 
negative. 

Gelatin stab: Liquefaction. 

Agar slant : Moderate, flat, faint yellow 
growth. 

Broth: Turbid in 5 days. 

Litmus milk : Acid. Peptonization 
after 16 days. 

Potato : Scant yellow growth with 
bleaching along line of growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Ammonia is produced. 

Acid from ghicose, maltose, lactose, 
sucrose, starch, glycerol and mannitol. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Source : Soil from Virginia, Louisiana 
and Missouri . 

Habitat: Soil. 

5a. Pseudomonas perlurida var. vir- 
giniana Kellerman et al. {loc. cit.). 
Does not grow on potato and liquefies 
gelatin rapidly. 

Source : Soil from Virginia. 

6. Pseudomonas iridescens Stanier. 
(Jour. Bact., 43, 1941, 542.) From Latin, 
iridescent. 

Rods: 0.2 to 0.3 by 1.5 to 7.0 microns, 
average length 5.0 to 6.0 microns, occur- 
ring singly. Non-motile. Gram-nega- 
tive. 



FAMILY PSEUDOMOXADACEAE 



175 



Sea water gelatin stab : Filiform growth. 
Liquefaction by some strains. 

Sea water agar colonies: Concave, 2 to 
3 mm in diameter, smooth, glistening, 
translucent, pale yellow, edge irregular. 
After 2 to 3 days a marked iridescence. 
Later colonies rough, opaque, bright 
yellow, sunken central portion with trans- 
lucent periphery. 

Sea water agar slant : Growth spreading, 
smooth, glistening, translucent, pale 
yellow, iridescent, butyrous. 

Sea water broth: Turbid, light yellow, 
granular pellicle. 

Indole not formed. 

Nitrites not produced from nitrates. 

Hydrogen sulfide not produced. 

Catalase positive. 

Urease negative. 

Acid from xylose, glucose, galactose, 
/actose, maltose, sucrose and cellobiose. 
No acid from arabinose. Starch and 
cellulose are attacked. 

Aerobic. 

Optimum temperature 23°C. Mini- 
mum o°C. Maximum 30°C. 

Salt range : 0.25 to 6.0 per cent. Opti- 
mum 1.0 to 4.0 per cent. 

Source : Sea water. 

Habitat : Common along the coast of 
the North Pacific. 

7. Pseudomonas turcosa (Zimmer- 
mann) Migula. {Bacillus turcosa Zim- 
mermann, Bakt. unserer Trink- und 
Nutzwasser, Chemnitz, 2, 1894, 32; 
Migula, Syst. d. Bakt. 2, 1900, 937; Flavo- 
bacteriiim turcosuvi Bergej^ et al., [Man- 
ual, 1st ed., 1923, 111.) From :\r. L. 
turcois, turquoise. 

Rods: 0.5 by 1.05 to 1.82 microns, oc- 
curring singly. A short polar fiagellum 
(Migula). Gram-negative. 

Gelatin colonies: Small, translucent, 
yellow. 

Gelatin stab: Small, yellow, convex 
surface growth, with slight brownish 
tint. Liquefaction, with grayish to 
greenish color in liquefied portion. 

Agar slant: Abundant, glistening, 
greenish to sulfur yellow streak. 



Broth: Slightly turbid with yellow 
sediment. 

Litmus milk: No coagulation. Yellow 
sediment. 

Potato : Clear chromium yellow growth 
over entire surface. 

Indole is not formed. 

Nitrites not produced from nitrates. 

Acid from glucose. Slight action on 
sucrose. 

Aerobic, facultative. 

Optimum temperature 30''C. 

Source : Isolated by Tataroff from a 
well in Dorpat (Die Dorpaten Wasser- 
bakterien, Inaug. Diss., 1891, 52, No. 24). 

Habitat : Water, sea water. 

8. Pseudomonas ochracea (Zimmer- 
mann) Chester. {Bacillus ochraceus 
Zimmermann, Bakt. unserer Trink- und 
Nutzwasser, Chemnitz, 1, 1890, 60; 
Chester, Determinative Bacteriology, 
1901, 316; Flavohacteritnn ochraceum Ber- 
gey et al., Manual, 1st ed., 1923, 110; 
Chromobacterium ochraceum Topley and 
Wilson, Princ. Bact. and Immun., 1, 
1931, 405.) From Greek, ochros, pale 
yellow. 

Rods : 0.7 to 0.8 by 1.2 to 4.5 microns, 
occurring in pairs and longer chains. 
Slow undulatory motion (Zimmermann). 
Polar flagella (Lehmann and Neumann, 
Bakt.Diag.,lAufl.,^, 1896,255). Gram- 
negative. 

Gelatin colonies : Pale yellow to golden, 
ochre yellow, slightly raised, with slightly 
fringed margin, granular. 

Gelatin stab : Yellowish to yellow-gray 
surface growth. Infundibuliform lique- 
faction. Pale yellow to ochre yellow 
sediment. 

Agar colonies: Thin, flat, j^ellowish, 
smooth. 

Agar slant: Thin, yellowish-gray to 
ochraceous growth. 

Broth: Slightly turbid, with pale yel- 
low sediment. 

Litmus milk: Medium becomes slimy; 
alkaline. 

Potato : Ochre-yellow streak. 

Indole is formed. 



176 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Nitrites not produced from nitrates. 
Hydrogen sulfide is formed. 
Aerobic, facultative. 
Optimum temperature 35°C. 
Source: Chemnitz tap water. 
Habitat: Water. 

9. Pseudomonas cerevisiae Fuhr- 
mann. (Fuhrmann, Cent. f. Bakt., II 
Abt., 16, 1906, 309; Flavohacterium 
cerevisiae Bergey et al., Manual, 1st ed., 
1923, 111.) From Latin, cerevisia, beer. 

Rods: Straight and slightly curved, 
0.6 by 1.5 to 2.0 microns, occurring singly 
and in chains. Motile, possessing tuft, 
four to six polar fiagella. Gram-negative . 

Gelatin colonies: Circular, white, 
slightly contoured, becoming brownish- 
yellow. 

Gelatin stab: Slight yellowish growth 
in stab. No liquefaction. 

Agar colonies: Thin, spreading, con- 
toured. 

Agar slant : Moist, glistening, thin, pale 
yellow, spreading, contoured. 

Litmus milk: Slow coagulation. 

Potato : Yellowish-brown, spreading 
growth. 

Indole not formed. 

Nitrites produced from nitrates. 

No gas from glucose . 

Aerobic, facultative. 

Optimum temperature 30°C. 

Source: Isolated from beer. 

Habitat: Unknown. 

10. Pseudomonas arguta McBeth. 
(McBeth, Soil Science, 1, 1916, 465; 
Cellulomonas arguata (sic) Bergey et al., 
Manual, 1st ed., 1923, 164.) From Latin, 
arguo, to show. 

Rods: 0.3 by 0.8 micron. Motile with 
one or two polar fiagella. Gram-negative. 

Gelatin stab: Moderate, yellowish 
growth. No liquefaction in 30 days. 

Agar colonies: Circular, slightly con- 
vex, soft, grayish-white, granular, entire. 

Agar slant: Scant, grayish-white 
growth. 

Potato agar slant: Moderate, yellow- 
ish, glistening. 



Broth: Turbid. 

Ammonia cellulose agar: Enzymatic 
zone 2 to 3 mm in 30 days. 

Filter paper broth : Paper is reduced 
to loose fiocculent mass which disinte- 
grates very readily on slight agitation. 
More rapid decomposition when the 
broth contains ammonium sulfate, po- 
tassium nitrate, peptone or casein as 
sources of nitrogen. 

Litmus milk : Acid, not digested. 

Potato: No growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Ammonia not produced. 

Acid from glucose, maltose, lactose, 
starch. No acid from glycerol, manni- 
tol or sucrose. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Source: Isolated twice from California 
soils. 

Habitat: Soil. 

1 1 . Pseudomonas subcreta McBeth 
and Scales. (McBeth and Scales, Bur. 
Plant Industry, U. S. Dept. Agr., Bui. 
266, 1913, 37; Cellulomonas subcreta 
Bergey et al., Manual, 1st ed., 1923, 164.) 
From Latin, sub, under, imperfect; creta 
chalk. 

Rods : 0.3 by 1.4 microns. Motile with 
one to five polar fiagella. Gram-negative. 

Gelatin stab : Filiform growth, no lique- 
faction. 

Cellulose agar: No surface growth. 
Moderate, generally faint yellow growth 
in medium, area of growth sunken. 

Agar slant: Glistening, smooth, moisl., 
vitreous to faint yellow. 

Starch agar : Enzymatic zone 2 to 4 mm. 

Broth : No growth. 

Litmus milk : No growth. 

Potato : Growth scanty, concave due to 
slight liquefaction, white to faint yellow. 
Bleached around growth. 

Indole not formed. 

Tr^ce of nitrites produced from ni- 
trates. 

Ammonia not produced. 

Acid from glucose, lactose, maltose, 



FAMILY PSEUDOMONADACEAE 



177 



sucrose and starch. No acid from glyc- 
erol or mannitol. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Habitat : Soil. 

12. Pseudomonas pictorum Gray 
and Thornton. (Gray and Thornton, 
Cent. f. Bakt., II Abt., 73, 1928, 89; 
Achromobacter pictorum Bergey et al., 
Manual, 3rd ed., 1930, 217.) From 
Latin, picti, the Pict.s of Eastern Scot- 
land. 

Rods: 0.5 to 0.8 by 1.5 to 5.0 microns. 
Motile usually with a single polar flagel- 
lum. Gram-negative. 

Gelatin colonies : Circular, greenish- 
yellow, convex, smooth, glistening, en- 
tire. 

Gelatin stab: Xo liquefaction. 

Agar colonies : Circular, j^ellow, convex, 
smooth, glistening, entire. 

Agar slant: Filiform, yellow, convex, 
smooth, glistening, entire. 

Broth: Turbid. 

Nitrites produced from nitrates. 

Starch not hydrolj^zed. 

Acid from glucose and maltose. 

Attacks phenol. 

Aerobic, facultative. 

Optimum temperature 25''C. 

Source: One culture from soil. 

Habitat : Soil. 

13. Pseudomonas lacunogenes Gor- 
esline. (Jour. Bact., 26, 1033, 447.) 
From Latin lacuno, dimple and gcnero, 
to produce. 

Short rods: 0.2 to 0.3 by 1.0 to 1.2 
microns, with pointed ends, occurring 
singly or in pairs. Motile with a single 
polar flagellum from 2 to 15 microns in 
length. Gram-negative. 

Plain gelatin stab : No growth. 

Nutrient gelatin stab : Growth brown- 
ish-yellow, half-way down stab, heavier 
at surface. No liquefaction. 

Nutrient agar colonies: Small, yellow; 
surface of the agar pitted or dimpled. 
After 5 days colonies 5 to 7 mm in diam- 



eter, orange-yellow, slightly raised, sur- 
rounded by a depression. 

Nutrient agar slant: Growth heavy, 
light orange-yellow ; consistency of warm 
butter; edge entire, slightly raised. 
Shallow depression formed on each side 
of streak . Agar softened beneath growth . 
Nutrient broth: Turbid in 48 hours. 
Light orange-yellow pellicle ; considerable 
viscous sediment. 

Litmus milk: Alkaline; butter-colored 
pellicle. Reduction in bottom of tube 
after 10 days. No curd. No digestion. 
Potato: Growth moderate, orange-yel- 
low, smooth. No darkening. 
Indole not formed. 
Nitrites not produced from nitrates. 
Starch agar plates not hydrolyzed. 
Utilizes arabinose, galactose, lactose, 
fructose, maltose, melezitose, raffinose, 
starch, xylose, glucose, mannose, su- 
crose, pectin, rhamnose, salicin and 
dextrin. No growth in dulcitol, erythri- 
tol, glycerol, sorbitol, mannitol or inulin. 
Limits of pH : 5.4 to 10.0.' 
Temperature relations : Optimum 28''C. 
Good growth at 25°C. Moderate growth 
at 20° and at 37°C. No growth at 10° 
and at 42°C. 
Facultative anaerobe. 
Distinctive characters: Softens agar; 
considerable change in viscosity of agar 
due to this digestion; utilization of 
ammonium sulfate as nitrogen source. 

Source : Three cultures isolated from 
an experimental trickling filter receiving 
creamery wastes. 

Habitat : Probably widely distributed 
in nature. 

14. Pseudomonas segnis Goresline. 
(Jour. Bact., 26, 1933, 452.) From Latin 
segne, non-energetic. 

Short rods: 0.2 to 0.3 by 1.0 to 1.2 mi- 
crons, with pointed ends, occurring singly 
or in pairs. Motile with a single polar 
flagellum. Gram-negative. 

Plain gelatin stab: No growth. 

Nutrient gelatin stab: Growth yellow, 
half-way down stab, best at surface. 
No liquefaction. 



178 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Nutrient agar colonies: Very small, 
light yellow; surface pitted. After 5 
days colonies 5 mm in diameter. 

Nutrient agar slant : Grovv'th heavy, 
orange-yellow, consistency of warm but- 
ter; edge entire, slightly raised; slight 
depression formed on each side of growth. 
Agar softened beneath growth. 

Nutrient broth: Turbid in 48 hours. 
No pellicle or surface growth. Moderate 
amount of sediment. Old cultures with a 
yellow ring at surface and occasionally a 
loose membrane. 

Litmus milk : Slightly alkaline after 10 
days. No reduction. No surface growth. 

Potato: Scant yellow-orange growth. 
No darkening. 

Indole not formed. 

Nitrites not produced from nitrates. 

No HoS produced. 

Starch not hydrolyzed. 

Utilizes arabinose, glucose, galactose, 
lactose, fructose, maltose, mannose, 
xylose, sucrose, melezitose and raffinose. 

Limits of pH : 5.8 to 9.0. 

Temperature relations : Optimum 28°C. 
Good growth at 25°C. Moderate growth 
at 20° and at 37°C. No growth at 10° 
and at 42°C. 

Facultative anaerobe. 

Distinctive characters: Softens agar; 
considerable change in viscosity of agar 
due to this digestion. 

Source : Isolated from an experimental 
trickling filter receiving creamery wastes. 

Habitat : Probably widely distributed 
in nature. 

15. Pseudomonas lemonnieri (Lasseur) 
comb. nov. (Bacillus lemonnieri Lasseur, 
Compt. rend. Soc. Biol. Paris, 74, 1913, 
47; Bui. de la Soc. des Sci. de Nancy, 
1924; Flavobacicrium lasseuri Bergey et 
al.. Manual, 3rd ed., 1930, 144.) Named 
for Prof. G. le Monnier, a French 
scientist. 

Rods: 0.5 to 0.7 by 1.0 to 2.0 microns, 
occurring singly and in pairs. Motile 



with a single polar flagellum. Gram- 
negative. 

Gelatin colonies (glucose) : Circular 
with blue center, a granular, yellow zone 
and a peripheral blue zone. Rapid 
liquefaction with blue crystals. 

Gelatin stab : Liquefied. 

Agar colonies: Circular, yellowish, 
lobate margin. 

Agar slant : Yellowish streak, smooth, 
glistening. 

Broth: Turbid with thin pellicle. 

Litmus milk: After 48 hours the sur- 
face of the milk becomes yellow to cream 
color turning blue. A soft coagulum is 
formed. 

Potato : Raised growth, Prussian blue 
in color, with variations. 

Indole is not formed. 

Nitrites produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 22° to 25°C. 

Habitat : Water. 

Appendix II:* The following inade- 
quately described species may belong 
to the genus Xanthomonas. 

Bacterium citri deliciosae Passalacqua. 
(Rev. Pat. Veg., 24, 1934, 27.) Isolated 
from Citrvs sp. 

Bacterium malvacearum var. harbadense 
Evelyn. (Ann. Rept. Agric. Barbados 
for 1926-27, 1928, 15.) Isolated from 
cotton. 

Pseudomonas amaranti (sic) Smith. (U. 
S. Dept. Agr., Div. Veg. Phys. and Path. 
Bull., 28, 1901, 153; Bacterium amaranthi 
Smith, Bact. in Relation to Plant Dis., 
3, 1914, 148; Phytomonas amaranthi 
Bergey et al., Manual, 1st ed., 1923, 186.) 
Isolated from diseased amaranthus. 
Growth in culture similar to Xantho- 
monas campestris and Xanthomonas hya- 
cinthi. 

Pseudomonas alutacea Migula. (Led- 
crgelber Bacillus, Tataroff, Die Dorpater 
Wasserbakterien, Inaug. Diss., Dorpat, 



* Prepared by Prof. Robert S. Breed, New York State Experiment Station, Geneva, 
New York, July, 1943. 



FAMILY PSEUDOMOXADACEAE 



179 



1891, 61 ; Migula, Syst. d. Bakt., 2, 1900, 
936.) Isolated from water. 

Pseudomonas graveolans Migula . ( Ba- 
cillus aquatilis graveolens Tataroff, Die 
Dorpater Wasserbakterien, Inaug. Diss., 
Dorpat, 1891, 48; Migula, Syst. d. Bakt., 
2, 1900, 934.) Isolated from water. 
Not Pseudomonas graveolens Levine and 
Anderson (Jour. Bact., 23, 1932, 343) 
isolated from musty eggs, and by Olsen 



and Hammer (Iowa State Coll. Jour. Sci., 
9, 1934, 125) from milk. 

Pseudomonas resinacea Migula. 
(Harzfarbener Bacillus, Tataroff, Die 
Dorpater Wasserbakterien, Inaug. Diss., 
Dorpat, 1891, 64; Migula, Syst. d. Bakt., 
2, 1900, 935.) Isolated from water. 

Xanthomonas taraxaci Niederhauser. 
(Phytopath., 53, 1943, 961.) Patho- 
genic on Russian dandelion (Taraxacum 
kok-saghz) . 



Genus III. Methanomonas Orla-Jensen.* 
(Cent, f . Bakt., II Abt., 22, 1909, 311.) 
Cells monotrichous, capable of obtaining energy from oxidation of methane to CO2 
and water. 
The type species is Methanomonas methanica (Sohngen) Orla-Jensen. 



1. Methanomonas methanica (Sohn- 
gen) Orla-Jensen. (Bacillus niethanicus 
Sohngen, Cent. f. Bakt., II Abt., 15, 
1906, 513; Orla-Jensen, Cent. f. Bakt., 
II Abt., 22, 1909, 311.) From methane. 

Short rods: 0.5 to 0.8 by 2.0 to 3.0 mi- 
crons, motile in young cultures by means 
of a single flagellum. In older cultures 
nearly spherical. Can be cultivated in 
an atmosphere composed of one part CH4 
and two parts air on washed agar contain- 
ing the necessarj^ inorganic salts. The 
growth is membranous. 



At the end of two weeks, the organisms 
changed an atmosphere containing 225 
ml. CH4 and 321 ml. O. to the follow- 



Clli ml. , 

CO2 78 ml. 

O2 172 ml. 

In addition, 21 ml. CO2 was dissolved 
in the liquid. 

Habitat : Presumably widely distrib- 
uted in soil. 



Genus IV. Acetobacter Beijerinck.j 
(Proc. Kon. Akad. v. Wetenschapp., Amsterdam, 2, 1900, 495.) 

Acetobacter aceti first appeared (Krai's Sammlung v. Mikroorg., Prague, 1898, 4) 
as a synonym of Bacterium aceti Hansen. Beijerinck (loc. cit.) mentions Acetobacter 
aceti in a footnote of a later paper. The genus name Acetobacter was accepted by 
Fuhrmann (Beiheft Bot. Centralbl., Orig., 19, 1905, 8) and others. From Latin, 
acetum, vinegar; baclrum, rod. 

Synonyms: ?Ulvina Kiitzing, Algae aquae dulcis, etc., 11th decade, 1837; Myco- 
derma Thompson, Ann. d. Chem. u. Pharmacie, 83, 1852, 89 ; ? Umbina Naegeli, Bericht 
liber die Verhandlingen der bot. Section der 33 Versammlung deutscher Xatur- 
forscher. und Arzter. Bot. Ztg., 1857, 760; Bacterium Lanzi, N. Giorn. bot. ital., 1876, 
257; Torula Saccardo, Atti Soc. Ven. Trent., 5, 1878, 315; Bacteriopsis (in part)Trevi- 
san, Atti Accad. Fisio-Medico-Statistica Milano, Ser. 4, 3, 1885, 103; Micrococcus 
Maggi, Jour. Microg., 10, ISSQ; Bacillus Schroeter, Kryptogamen Flora von Schlesien, 
3, 1, 1886, 161; Termobacterium Zeidler, Cent. f. Bakt., II Abt., 2, 1896, 739; Acetobac- 

* Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, December, 1922. 

t Revised by Dr. C. D. Kelly, McGill Univ., Montreal, P. Q., Canada, July, 1938; 
further revision by Dr. Reese H. Vaughn, Univ. of California, Berkeley, California 
June, 1943. 



180 MANUAL OF DETERMINATIVE BACTERIOLOGY 

terium Ludwig, in abstract of Hoyer's Inaug. Diss., Cent. f. Bakt., II Abt., 4, 1898, 
867; Acetimonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 312. 

In addition, the sub-generic names E uacetobacter and Acetogluconobacier have 
been proposed by Asai, Jour. Agr. Soc. Japan, 11, 1935, 502. Tlie genus Gluconobacter 
and the sub-genera Eugluconobacter and Gluconoacetobacter Asai {loc. cit.) may be 
synonyms in whole or in part. 

Individual cells ellipsoidal to long and rod-shaped, occurring singly, in pairs, or 
in short or long chains. Motile with polar flagella, or non-motile. Involution forms 
may be spherical, elongated, filamentous, club-shaped, swollen, curved or even 
branched. Young cells Gram-negative; old cells often Gram-variable. Obligate 
aerobes; as a rule strongly catalase positive, sometimes weakly so. Oxidize various 
organic compounds to organic acids and other oxidation products which may undergo 
further oxidation. Common oxidation products include acetic acid from ethyl alcohol, 
gluconic and sometimes ketogluconic acid from glucose, dihydroxyacetone from gly- 
cerol, sorbose from sorbitol, etc. Nutritional requirements vary from simple to com- 
plex. Development generally best in yeast infusion or yeast autolysate media with 
added ethyl alcohol or other oxidizable substrate. Optimum temperature variable 
with the species. Widely distributed in nature where they are particularly abun- 
dant in plant materials undergoing alcoholic fermentation; of importance to man for 
their role in the completion of the carbon cycle and for the production of vinegar. 

The type species is Acetobacter aceti (Kiitzing) Beijerinck. 

Key to species of genus Acetobacter. 
I. Oxidize acetic acid to carbon dioxide and water. 

A. Capable of utilizing ammonium salts as a sole source of nitrogen (Hoyer's 

solution).* 

1. Acetobacter aceti. 

B. Do not utilize ammonium salts as a sole source of nitrogen.* 

1. Forms a thick, zoogloeal, cellulose membrane on the surface of liquid 

media. 

2. Acetobacter xylinum. 

2. Do not form a thick, zoogloeal membrane on the surface of liquid 

media. 

3. Acetobacter rancens. 

3a. Acetobacter pasteurianum. 
3b. Acetobacter kuetzingianum . 
II. Do not oxidize acetic acid. 

A. Form pigments in glucose media. 

1. Dark brown to blackish pigment. 

4. Acetobacter melanogenum. 

2. Pink to rose pigment. 

5. Acetobacter roseinn. 

B. Do not form pigments. 

1. Optimum temperature 30° to 35°C. 

6. Acetobacter suboxydans. 

2. Optimum temperature 20° to 25°C. 

7. Acetobacter oxydans. 

* It is not known with certainty whether Acetobacter pasteurianum and Acetobacter 
kuetzingianum are capable of using inorganic nitrogen as a sole source of nitrogen 
for growth. However, since these two species are among those first described it is 
advisable to retain them for the present. See Acetobacter rancens Beijerinck. 



FAMILY PSEUDOMOXADACEAE 



181 



1. Acetobacter aceti (Kiitzing) Beijer- 
inck. (Ulvina aceti Kiitzing, Algae 
aquae dulcis etc., Uth decade, 1837; 
Mycoderma aceti Thompson, Ann. d. 
Chem. u. Pharmacie, 83, 1852, 89; Um- 
bina aceti Naegeli, Bericht iiber die 
Verhandlingen der bot. Section der 33 
Versammlung deutscher Naturforscher 
und Arzter. Bot. Ztg., 1857, 760; Bac- 
terium aceti Lanzi, N. Giorn. bot. ital., 
1876, 257; Torula aceti Saccardo, Atti 
Soc. Ven. Trent., 5, 1878, 315; Bacteriop- 
sis aceti Trevisan, Atti della Accademia 
Fisio-Medico-Statistica in Milano, Ser. 
4, 3, 1885, 103; Micrococcus aceti Maggi, 
Jour. Microg., 10, 1886; Bacillus aceti 
Schroeter, Kryptogamen Flora von Schle- 
sien, 3, 1, 1886, 161; Bacillus aceticus 
Fliigge, Die Mikroorganismen, 1886, 313; 
Beijerinck, Krai's Sammlung v. Mikro- 
org., Prague, 1898, 7; Beijerinck, Proc. 
Kon. Akad. v. Wetensch., Amsterdam, 
2, 1900, 495; Bacterium hansenianum 
Chester, Man. Determ. Bact., 1901, 126.) 
From Latin acetum, vinegar. 

Rods: 0.4 to 0.8 by 1.0 to 2.0 microns, 
occurring singly and in long chains, fre- 
quently showing large club-shaped forms. 
Stain yellow with iodine solution. Mo- 
tility variable. Motile cells possess a 
single polar flagellum (Vaughn, Jour. 
Bact., 46, 1943, 394). Forms large, 
shiny colonies on beer gelatin containing 
10 per cent sucrose. 

Forms slimy pellicle on fluid media, or 
ring or turbidity without pellicle. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid from 
arabinose, fructose, galactose, sorbose, 
sucrose, maltose, lactose, raffinose, dex- 
trin, starch, glycogen, inulin, methyl 
alcohol, isopropyl alcohol, butyl alcohol, 
isobutyl alcohol, amyl alcohol, glycerol, 
erythritol, mannitol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Es- 
sigind., B, 1898, 147). 
Aerobic. 

Distinctive characters : Marked oxida- 
tive power causing rapid and complete 
oxidation of substrate as glucose or ethyl 
alcohol; ability to utilize inorganic nitro- 



gen salts as a sole source of nitrogen 
(Hoj^er, Inaug. Diss., Leiden, 1898, 43; 
Beijerinck, Cent. f. Bakt., II Abt., 4, 
1898, 215) ; growth and oxidative activity 
in association with fermenting yeasts 
(Vaughn, Jour. Bact., 36, 1938, 360). 

Optimum temperature 30°C. Growth 
occurs between 10° and 42°C. 

Habitat: Vinegar; souring fruits, vege- 
tables and beverages. 

2. Acetobacter xylinum (Brown) Hol- 
land. (Bacterium xylinum Brown, Jour. 
Chem. Soc, London, 49, 1886, 439; 
Holland, Jour. Bact., 5, 1920, 216; Ba- 
cillus xylinus Holland, ibid., 221.) 
From Gr. xijlinus, wooden (in reference 
to the cellulose in the membrane). 

Rods, about 2 microns long, occurring 
singly and in chains. The cells have a 
slimy envelope which gives the cellulose 
reaction. 

A film forms on the surface of liquids. 
This film becomes cartilagenous and falls 
to the bottom. This zoogloeal film forms 
on all liquid media in which growth 
occurs; the nature of the medium influ- 
ences the thickness of the film which may 
vary from 1 to 250 millimeters. 

X-ray pattern studies made by Khou- 
vine, Champetier and Sutra (Compt. 
rend. Acad. Sci. Paris, 194, 1932, 208) 
and by Barsha and Hibbert (Can. Jour. 
Research, 10, 1934, 170) have shown that 
the cellulose contained in the membranes 
formed by Acetobacter xylinum is identi- 
cal with cotton cellulose. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid from 
arabinose, fructose, galactose, maltose, 
lactose, raffinose, dextrin, starch, methyl 
alcohol, isopropyl alcohol, butyl alcohol, 
isobutyl alcohol, amyl alcohol, manni- 
tol and acetaldehyde (Henneberg, Die 
deutsch. Essigind., 2, 1898, 147). 
Aerobic. 

Distinctive character: The production 
of thick, leathery, zoogloeal cellulosic 
membranes on the surface of liquids. 
Optimum temperature 28°C. 



182 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Habitat : Vinegar; souring fruits, vege- 
tables and beverages. 

3. Acetobacter rancens Beijerinck. 
{Bacterium rancens Beijerinck, Cent. f. 
Bakt., II Abt., 4, 1898, 211; Beijerinck, 
Krai's Sammlung v. Microorg., Prague, 
1898, 4.) From L. rancens, being rancid. 

Beijerinck {loc. cit.) in a footnote 
stated that "two of the many varieties of 
B. rancens have been described by Hen- 
neberg under the names of B. oxydans 
and B. acetosum. Hansen erroneously 
called this species B. aceti as did Brown. 
Neither Hansen nor Brown knew B. 
aceii Pasteur." No further morphologi- 
cal description is given. 

The following description is taken in 
part from a study of a culture of Aceto- 
bacter rancens received from Kluyver 
(Vaughn). 

Rods with the usual morphological ap- 
pearance of cultures of acetic acid 
bacteria. Gram-negative. Motility 
variable. Motile cells possess a single 
polar flagellum (Vaughn, Jour. Bact., 
46, 1943, 394). Involution forms com- 
monly appear as filaments and enlarged 
cells. 

Wort agar slant: Growth abundant, 
butyrous, pale-buff in color in one week. 

Yeast infusion, glucose, calcium 
carbonate slant : Growth abundant, bu- 
tyrous and cream-colored in one week. 

With petri dish cultures well isolated 
colonies are large, smooth and butyrous 
on either medium. 

Broth cultures containing peptone or 
yeast infusion form a mucilaginous, slimj^ 
pellicle. Beijerinck {loc. cit.) called 
this polysaccharide pellicle, cellulose-like 
and intimated that the mucilaginous 
material in the pellicle was somewhat 
different from that produced by Aceto- 
bacter xylinum. The pellicle material 
stained blue when treated with iodine 
and hydroiodic acid. 

Acid from glucose, ethyl alcohol, propyl 
alcohol, butyl alcohol, glycol, adonitol, 
mannitol and sorbitol. No acid from 
numerous other compounds tested. 



Distinctive character : Production of a 
thin, mucilaginous, slimy, polysaccharide 
membrane on the surface of liquids as 
compared with the tMck, true cellulose 
membrane of Acetobacter xylinum grown 
under the same conditions. Beijerinck 
{loc. cit.) reported the production of a 
cellulose-like membrane with some cul- 
tures of Acetobacter rancens. 

Source : Isolated from shavings in the 
quick vinegar process. 

Habitat : Found in fermented grain 
mash, malt beverages, mother of vinegar. 

Beijerinck (Cent. f. Bakt., II Abt., 4, 
1898, 211) thought that the next two 
species were hardly more than varieties 
of Acetobacter rancens. 

3a. Acetobacter pasteurianum (Han- 
sen) Beijerinck. {Mycoderma ■pasteuri- 
anum Hansen, Compt. rend. d. Trav. d. 
Lab. d. Carlsberg, 1, 1879, 96; Bacteriuvi 
pasteurianum Zopf, Die Spaltpilze, 2 
Aufl., 1884, 49; Beijerinck, Krai's Samm- 
lung V. Microorg., Prague, 1898, 7.) 
Named for Pasteur, the French chemist 
and bacteriologist. 

Rods: 0.4 to 0.8 by 1.0 micron, 
occurring singly and in chains, at times 
showing thick, club-shaped forms. Mo- 
tility variable. Motile cells possess a 
single polar flagellum (Vaughn, Jour. 
Bact., 46, 1943, 394). Stains blue with 
iodine. 

Wort gelatin colonies: Small, circular, 
entire, gray, slimy. 

Forms a dry, wrinkled folded pellicle 
on double beer with one per cent alcohol. 

Meat infusion gelatin: Widespread, 
later rosette form, toothed. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid from 
arabinose, fructose, galactose, sorbose, 
sucrose, maltose, lactose, raffinose, 
dextrin, starch, glycogen, inulin, methyl 
alcohol, isopropyl alcohol, butyl alcohol, 
isobutyl alcohol, amj'l alcohol, glycerol, 
erythritol, mannitol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind., 2, 1898, 147). 

Aerobic. 



FAMILY PSEUDOMONADACEAE 



183 



Optimum temperature 30°C. Growth 
occurs between 5° and 42°C. 

Habitat : Vinegar; beer and beer wort. 

3b. Acetobacter kuetzingianum (Han- 
sen) Bergey et al. (Bacterium kuetzingi- 
anum Hansen, Compt. rend. d. Trav. d. 
Lab. d. Carlsberg, 3, 1894. 191; Bergey 
et al., Manual, 1st cd., 1923, 35.) Xamed 
for Kuetzing, the German botanist. 

Short, thick rods, occurring singly. 
Rarely forming chains of notable length. 
Capsule stained blue with iodine and witli 
potassium iodide. Xon-motile. 

Double beer gelatin colonies : Small, 
entire, with vermiform surface. 

Wort gelatin colonies: Small, entire, 
with surface free of wrinkles. 

Double beer: Forms a rather thick, 
folded pellicle. Distinguished from A ce- 
tobacter aceti in showing heavier growth 
above the surface of the media. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid from 
arabinose, fructose, galactose, sorbose, 
sucrose, maltose, lactose, raffinose, dex- 
trin, starch, glycogen, inulin, methyl 
alcohol, isopropyl alcohol, butyl alcohol, 
isobutyl alcohol, amyl alcohol, glycerol, 
erythritol, mannitol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind., £, 1898, 147). 
Aerobic. 

Optimum temperature 34°C, maximum 
42°C, minimum 6 to 7°C. 
Habitat : Beer. Found in double beer. 



gelatin becomes insoluble in boiling water 
and in trypsin solution. 

Beer- or wort-gelatin plates: Charac- 
teristic dark brown, wide-spreading, 
diffuse areas. 

Tap water-agar-glucose-peptone-po- 
tassium phosphate-iron citrate-chalk 
medium: In 24 hours at 30°C, black, 
spreading, diffuse areas. 

Utilizes peptone as a source of nitrogen. 
Produces the pigment from peptone only 
if maltose or glucose is present as a 
source of carbon. When grown in glu- 
cose-peptone broth with CaCOs at 25° 
to 30°C, black pigment is produced after 
several weeks, and the carbonate is 
changed to calcium gluconate. 

Pigment : The pigment causing the 
brown coloration is an aromatic substance 
which is blackened by iron salts. Re- 
duces alkaline solutions of silver and 
mercury, blackening them. 

Oxidizes mannitol and sorbitol to fruc- 
tose and sorbose. Does not attack sucrose 
and fructose. ]\Iuch gluconic acid is 
produced. Acid from glucose and mal- 
tose. Acetic acid produced from alcohol. 
Distinctive character: The formation 
of dark brown to black pigment in media 
containing a suitable substrate; par- 
ticularly glucose. 
Source : Isolated from beer. 
Habitat: Causes light-colored beer to 
become darker brown. It is a very strong 
beer-vinegar bacterium. Also found in 
souring fruits. 



4. Acetobacter melanogenum Bei- 
jerinck. (Cent. f. Bakt., II Abt., 29, 
1911, 175.) From Greek melas (mela7i), 
black; -genes, producing. 

Rods : Xon-motile or motile. Motile 
cells possess a single polar flagellum 
(Vaughn. Jour. Bact., 46, 1943, 394). 

Gelatin: Apparent liquefaction prob- 
ably caused by acid, not an enzyme. 
When hold on artificial media for some 
time, the power of liquefying gelatin is 
lost, probably due to a slower production 
of acid. Deep brown pigment produced ; 



5. Acetobacter roseum Vaughn. 
{Bacterium hoshigaki var. rosea Taka- 
hashi and Asai, Cent. f. Bakt., II Abt., 
82, 1930, 390; Acetobacter fioshigaki 
Bergey et al., :Manual, 4th ed., 1934, 39; 
Vaughn, Wallerstein Lab. Communica- 
tions, o, Xo. 14, 1942, 20.) From Latin, 
rasa, rose. 

Rods : 0.7 to 0.9 by 1.5 to 1.8 microns, 
generally occurring singly, at most in 
pairs, often in chains. Xon-motile. 
Pellicle on fluid media yields no starch 
or cellulose reaction. 



184 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Koji (a mixture of rice and mold spores 
used to start fermentation of Japanese 
bread and sak6) extract agar colonies : 
Small, granular, circular, glistening, 
umbonate, becoming brownish. 

Wort agar colonies : Circular, milky- 
white, becoming brownish in center and 
yellowish at periphery. 

Glucose sake agar: Circular, milky- 
white, granular, umbonate, entire. 

Hoshigaki (dried persimmons) extract 
agar: Circular, milky -white, granular, 
becoming yellowish-brown in the center 
and grayish-white at the periphery. 

Koji extract agar streak : Grayish- 
white, glistening with ciliate margin, 
becoming purple brown to brown. 

Koji extract: Turbid with thin film, 
ascending on wall of tube. 

Bouillon: Turbid with ring formation. 

Yeast infusion glucose agar : Colonies 
similar to those on wort agar. 

Yeast infusion glucose broth : Turbid 
with thin, ascending film. 

Red color produced on sake wort agar 
and all media containing calcium 
carbonate. 

Acid from glucose, fructose, galactose, 
arabinose, glycerol, mannitol, ethyl and 
propyl alcohol. No acid from maltose, 
sucrose, lactose, raflinose, dextrin, starch, 
inulin, sorbitol, glycogen, isodulcitol and 
methyl alcohol. 

Forms gluconic acid from glucose. 

Aerobic. 

Optimum temperatures 30° to 35°C; 
maximum 40° to 41°C; minimum 10° 
to 15°C. 

Thermal death point 50°C for 5 
minutes. 

Distinctive character : The formation 
of a rose to red pigment in suitable media ; 
particularly those containing glucose 
and calcium carbonate. 

Source : Isolated from fermenting mash 
of dried persimmons (hoshigaki), and 
souring figs and dates. 

Note: Vaughn, Wallerstein Lab. Com- 
munications, 5, No. 14, 1942, 20, has 
proposed the name Acetohacter roseiun to 
replace the name Acetohacter hoshigaki. 



As originally described, this organism 
was given the name Bacterium hoshigaki 
var. rosea by Takahashi and Asai (loc. 
cit.) without the authors having first 
named and described the species Bac- 
terium hoshigaki. The Japanese word 
"hoshigaki" has been used in a confus- 
ing manner viz. Takahashi and Asai, 
loc. cit. {Bacterium industrium var. 
hoshigaki) and Takahashi and Asai, 
Jour. Agr. Chem. Soc. Japan, 9, 1933, 351 
and Cent. f. Bakt., II Abt., 87, 1933, 385 
{Bacteriuin hoshigaki var. glucuronicum 
I, II and III). None of these Japanese 
names are in the form of true binomials. 

6. Acetobacter suboxydans Kluyver 
and de Leeuw. (Paper read at the con- 
vention of the Dutch Society of Microbiol- 
ogy, Utrecht, December, 1923, see Tijd- 
schrift V. Vergelijkende Geneeskunde, 
10, Afl. 2-3, 1924.) From L. sub, under, 
less; Gr. oxys, sharp, acid; dans, giving, 
i.e. less acid giving; less oxidizing. 

Short rods : Occurring singly or in 
chains. Non-motile. Morphologically 
like Acetobacter rancens. 

Forms very thin, hardly visible pellicle 
on fluid media. 

Wort agar colonies: Very small, circu- 
lar, slightly yellow. 

Acid from ethyl alcohol, propyl alco- 
hol, glycol, glucose, glycerol and sorbitol. 

Optimum temperature 30°C. 

Distinctive character: Partial oxida- 
tion of substrates as indicated by the 
formation of calcium 5-keto gluconate 
crystals on the surface of agar slants con- 
taining glucose and calcium carbonate. 

Source : Isolated from spoiled beer. 

Habitat : Beer. 

7. Acetobacter oxydans (Henneberg) 
Bergey et al. (Bacteritmi oxydans Hen- 
neberg, Cent. f. Bakt., II Abt., 3, 1897, 
223; Bacillus oxydans Migula, Syst. d. 
Bakt., 2, 1900, 800; Bergey et al.. Manual, 
1st ed., 1923, 36.) From Gr. oxys, sharp, 
acid; dans, giving. 

Rods: 0.8 to 1.2 by 2.4 to 2.7 microns, 
occurring singly and in chains. Motile 



FAMILY PSEUDOMONADACEAE 



185 



cells possess a single polar flagellum 
(Vaughn, Jour. Bact., ^6, 1943, 394). The 
chains show bud-like swellings. 

Gelatin colonies : Circular, becoming ir- 
regular in shape with peculiar ramifica- 
tions. 

Acid from arabinose, fructose, glucose, 
galactose, sucrose, maltose, raffinose, 
dextrin, ethyl alcohol, propyl alcohol, 
erythritol, mannitol, glycol and glycerol. 
No acid from sorbose, lactose, starch, 
glycogen, inulin, methyl alcohol, isopro- 
pyl alcohol, butyl alcohol, isobutyl 
alcohol, amyl alcohol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind.,;2, 1898, 147). 

Aerobic. 

Optimum temperature 18° to 21°C. 

Distinctive characters: Low optimum 
temperature for growth and oxidation of 
substrates; and the ability to oxidize a 
large number of substrates. 

Habitat: Beer. 



Appendix: The following species have 
been described, but until more compara- 
tive studies have been made, no change 
in nomenclature is recommended or 
advisable. 

1. Acetobacter zeidleri Beijerinck. 
(Termobaderium aceti Zeidler, Cent. f. 
Bakt., II Abt., 2, 1896, 739; Bacterium 
zeidleri Beijerinck, Acetobacter zeidleri 
Beijerinck, Krai's Sammlung v. Mikro- 
org., Prague, 1898, 7; Bacillus zeidleri 
Migula, Syst. d. Bakt., 2, 1900, 801; 
Acetobacter lindneri Bergey et al., Man- 
ual, 1st ed., 1923, 36.) Named for A. 
Zeidler, who first isolated this species. 

Rods, occurring singly and in chains, 
showing large sausage-shaped involution 
forms. Motile with a single polar flagel- 
lum (Zeidler, Cent. f. Bakt., II Abt., 4, 
1898, 669). 

Wort gelatin: Small, circular, slightly 
granular, yellowish-brown, entire 
colonies. No liquefaction. 

Dirty, yellowish-brown pellicle on 
liquid media. 

Wort gelatin slant : Strongly glistening, 



transparent, whitish in center, smooth, 
very weakly liquefied. 

Potato: Very scant growth. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid from 
arabinose, fructose, galactose, maltose, 
lactose, rafhnose, dextrin, glycogen 
methyl alcohol, isopropyl alcohol, butyl 
alcohol, isobutyl alcohol, amyl alcohol, 
glycerol, mannitol and acetaldehyde 
(Henneberg, Die deutsch. Essigind., 2, 
1898, 147). 

Aerobic, facultative. 

Optimum temperature 25°C. 

Habitat: Beer wort. 

2. Acetobacter acetosum (Henneberg) 
Bergey et al. {Bacterium acetosutn Hen- 
neberg, Cent. f. Bakt., II Abt., 3, 1897, 
223; Bergey et al.. Manual, 1st ed., 1923, 
36.) From Latin, acelum, vinegar. 

Rods : 0.4 to 0.8 by 1 .0 micron, occurring 
singly and in chains. Non-motile. 
Stains yellow with iodine. 

On beer, yeast water and glucose solu- 
tions a firm, coherent, uniform, smooth, 
white film that becomes folded (Henne- 
berg, Garungsbakt., 2, 1926, 201). 

Acid from glucose, galactose, ethyl 
alcohol, and propyl alcohol. No acid 
from arabinose, fructose, sorbose, sucrose, 
maltose, lactose, raffinose, dextrin, 
starch, glycogen, inulin, methyl alcohol, 
isopropyl alcohol, butyl alcohol, iso- 
butyl alcohol, amyl alcohol, glycerol, 
erythritol, mannitol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind., 2, 1898, 147). 

Optimum temperature 28°C, maximum 
36°C, minimum 8°C (Henneberg, Cent, 
f. Bakt., II Abt., 4, 1898, 14). 

Habitat : Beer. 

3. Acetobacter ascendens (Henne- 
berg) Bergey et al. {Bacterium ascen- 
dens Henneberg, Zeitschr. f. deutsche 
Essigind., Berlin, No. 19 to 23, 1898, 145; 
also see Cent. f. Bakt., II Abt., 4, 1898, 
933; Bergey et al., Manual, 1st ed., 1923, 
37.) From Latin, ascendo, pp. ascen- 
dens, ascending. 

Rods, occurring singly, rarely in chains. 



186 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Non-motile. Do not give the cellulose 
reaction with iodine solution. 

Glucose gelatin colonies : Dry, white, 
with white area surrounding the colony. 

Fluid cultures have a tough pellicle 
rising on the wall of the flask. 

Acid from ethyl alcohol, propyl alcohol 
and glycol. No acid from arabinose, 
fructose, glucose, galactose, sucrose, 
maltose, lactose, raffinose, dextrin, 
starch, methyl alcohol, isopropyl alcohol, 
butyl alcohol, isobutyl alcohol, amyl 
alcohol, glycerin, mannitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind., 2, 1898, 147). 

Aerobic. 

Optimum temperature 31°C. 

Habitat: Isolated from vinegar and 
from red wine. 

4. Acetobacter plicatum Fuhrmann. 
(Beihefte z. bot. Centralbl., Orig., 19, 
1905, 8.) Description given in Cent. f. 
Bakt., II Abt., 15, 1906, 377. From 
plicatus, folded. 

Rods : 0.55 to 0.7 by 0.75 to 0.9 microns 
when grown on agar at 28° to 30°C. 
Young streak cultures 0.4 to 0.6 by 1.4 
to 1.6 microns with homogeneous staining 
when grown on beef-extract-gelatin at 
22°C. 0.5 by 1.5 to 1.7 microns with 
uneven staining (polar) when grown on 
wine gelatin. At about 40°C the organ- 
isms form swollen and greatly elongated 
forms. Non-motile. 

Agar slant : Pale yellowish, translucent 
growth. 

Alcohol-free beer with glucose and 
sucrose: Turbid with thick pellicles. 

Potato : Growth limited. 

Ferments alcohol to form acetic acid. 

Optimum temperature 28° to 30°C. 

Habitat: Wine. 

5. Acetobacter acetigentmi (Henne- 
berg) Bergey et al. {Bacterium aceti- 
genum Henneberg, Cent. f. Bakt., II 
Abt., 4, 1898, 14; Bacillus aceligenuni 
Migula, Syst. d. Bakt., 2, 1900, 801; 
Bergey et al., Manual, 1st ed., 1923, 
35.) From Latin, producing vinegar. 



Rods, occurring singly and in pairs. 
0.8 to 1.2 by 1.2 to 1.4 microns. Motile. 
Cells give a cellulose reaction with H0SO4 
and iodine. 

Glucose gelatin colonies : Raised, gray- 
ish, slimy. 

Fluid cultures show a tough, slimy 
pellicle. 

Acid from glucose, ethyl alcohol, 
propyl alcohol and glycol. No acid 
from arabinose, fructose, galactose, sor- 
bose, sucrose, maltose, lactose, raffinose, 
dextrin, starch, glycogen, inulin, methyl 
alcohol, isopropyl alcohol, butyl alcohol, 
isobutjd alcohol, amyl alcohol, glycerol, 
erythritol, mannitol, dulcitol and acetal- 
dehyde (Henneberg, Die deutsch. Essig- 
ind.,;2, 1898, 147).' 

Aerobic. 

Optimum temperature 33°C. Thermal 
death point 43° to 45°C for 5 minutes. 

Habitat : Vinegar. 

6. Acetobacter industrium (Henne- 
berg) Bergey et al. {Bacterium indus- 
trium Henneberg, Zeitschr. f. deutsche 
Essigindustrie, Berlin, 1898; Cent. f. 
Bakt., II Abt., 4, 1898, 933; Bacillus 
industrius Migula, Syst. d. Bakt., 2, 
1900, 801; Bergey et al.. Manual, 1st ed., 
1923, 36.) From Latin industrius, 
diligent. 

Rods: 0.3 to 0.8 by 2.4 to 20 microns, 
occurring singly and in chains. No 
distinct color produced with iodine. 
Motile. 

Forms pellicle on fluid culture media. 

Acid from arabinose, fructose, glucose, 
galactose, sucrose, maltose, lactose, raffi- 
nose, starch, dextrin, ethyl alcohol, 
propyl alcohol, glycol, glycerol and 
mannitol. No acid from isopropyl alco- 
hol, butjd alcohol, isobutyl alcohol, amyl 
alcohol and acetaldehyde (Henneberg, 
Die deutsch. Essigind., 2, 1898, 147). 

Aerobic. 

Optimum temperature 23°C. Maxi 
mum 35°C. Minimum 8°C. 

Habitat : Beer wort. 



FA&ULY PSEUDOMONADACEAE 



187 



7. Bacterium schuezenbachii Henne- 
berg. (Die deutsche Essigind., No. 
11-18, 1906; also Cent. f. Bakt., II Abt., 
17, 1906, 790.) Named for Schiizenbach, 
the inventor of the German quick vinegar 
process. 

Rods: 0.3 to 0.4 bj^ 1.0 to 3.6 microns, 
occurring singly, in pairs and chains. 
The cells are round, oval or elongated, not 
infrequently sickle-shaped or irregularly 
bent with rounded or pointed ends. 
Not stained with iodine. Non-motile. 

Wort gelatin colonies : Round, shin}', 
transparent with yellowish-brown cen- 
ters. 

A non-coherent fihn produced on the 
surface of liquid media. 

Acid from arabinose, fructose, glucose, 
galactose, maltose, lactose, dextrin, ethyl 
alcohol, propyl alcohol, glycerol and 
erythritol. Small amount of acid from 
sucrose and raffinose. No acid from 
mannitol (Henneberg, Handbuch d. Gar- 
ungsbakt., 2, 1925, 239). 

Temperature relations : Optimum 25° 
to 27.5^0. Scant growth at 34° to 35"C 
and 13° to 15°C. Xo growth at 37° and 
7.5°C. 

Source: Isolated from vinegar in the 
quick vinegar process. 

Habitat: Produces acetic acid in quick 
vinegar process. 

8. Bacterium xylinoides Henneberg. 
(Die deutsche Essigind., No. 11 to 18, 
1906; also Cent. f. Bakt., II Abt., 17, 
1906, 794.) From Greek, woody. 

Rods: 0.5 to 0.8 microns (round cells) 
and 0.5 to 1.2 microns (long forms), oc- 
curring singly, in pairs or chains, cells 
round and as short and long rods. The 
thick membrane like that produced by 
Acetohactcr xylinum gives the reaction 
for cellulose with iodine and sulfuric 
acid, but the thin membrane does not. 

Wort gelatin : Colonies are produced 
like drops of water, often with light brown 
kernels in the center. 

Wort gelatin streak : Growth transpar- 
ent at first, later whitish. 



Three types of membrane on fluid 
media are formed by this species and all 
three may even be found on a culture at 
one time. A thin, firm, coherent mem- 
brane like that of Bacterium orleanense 
may be formed or one that is voluminous, 
scumlike (like coagulated egg-white), 
coherent, somewhat slimj'^ and finally 
thick like that of Bacterium xylinum. 
Also a membrane may be formed that is 
intermediate in type. 

Acid from arabinose, glucose, galac- 
tose, sucrose, maltose, ethyl alcohol, 
propyl alcohol, glycerol and erythritol. 
Small amount of acid from fructose and 
mannitol. 

Temperature relations : Optimum 28°C. 
Slight growth at 14°C. No growth at 
6°C. 

Source : Isolated from wine vinegar 
from the Rhine and other sections. 

Habitat : Found in vinegar made by the 
Orleans method. 

9. Bacterium orleanense Henneberg. 
(Die deutsche Essigind., No. 11-18, 1906; 
also Cent. f. Bakt., II Abt., 17, 1906,792.) 
Latinized, of Orleans. 

Rods: 0.4 to 0.5 by 1.2 to 2.1 microns, 
occurring singly or in chains. The cells 
are round, elongated or as involution 
forms, with straight or curved cells 
appearing. Not stained with iodine. 
Non-motile. 

Wort gelatin : Colonies irregular in 
form, whitish in color, about 1 mm. in 
diameter in 2 days. 

Wort gelatin streak: Growth often 
slimy, transparent, licjuid mass with 
yellowish-brown sediment. 

Wort agar streak: Some strains form 
isolated, moist, slimy, transparent colo- 
nies and on the water of condensation 
isolated whitish colonies are formed. 
Other strains form a coherent, trans- 
parent coating with a light brown pre- 
cipitate later and individual, distinct, 
round colonies of the same color. 

Characteristic of this species is the firm 
coherent film on the surface of liquid 
media. 



188 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Acid from arabinose, glucose, galac- 
tose, maltose, lactose, raffinose, dextrin, 
ethyl alcohol, propyl alcohol, glycerol, 
erythritol and mannitol. Small amount 
of acid from fructose and sucrose (Hen- 
neberg, Handbuch d. Garungsbakt., 2, 
1926,239). 

Temperature relations : Optimum 20° 
to 30°C. Slight growth at 35° to 36°C 
and 14° to 15°C. No growth at 39° and 
at 7° to 8°C. 

Source : Isolated from vinegar in the 
quick vinegar process. 

Habitat: Can be used both in the 
quick or German process and the Orleans 
method of making vinegar. 

10. Bacterium vini acetati Henneberg. 
(Die deutsche Essigind., No. 11-18, 1906; 
also Cent. f. Bakt., II Abt., 17, 1906, 
797.) From Latin vinum, wine and 
acetum, vinegar. 

Rods: 0.3 to 0.8 by 0.8 to 2.0 microns, 
occurring singly, in pairs and sometimes 
as short chains of three; cell round, oval 
or slightly elongated, and rarely moder- 
ately long forms. Streptococcus-like 
cells are found on older agar cultures and 
spindle forms in beer gelatin with 10 per 
cent sucrose. 

Wort gelatin : Round, moist, shiny, 
transparent colonies with whitish sedi- 
ment in the center. 

The film on liquid media is not strongly 
coherent and the liquid is cloudy. 

Acid from arabinose, fructose, glucose, 
galactose, sucrose, maltose, raffinose, 
dextrin, ethyl alcohol, propyl alcohol, 
glycerol and erythritol. No acid from 
lactose (Henneberg, Handbuch d. Gar- 
ungsbakt., .?, 1925, 239). 

Optimum temperature 28° to 33°C. 

Source: Wine vinegar. 

Habitat : Found in vinegar made by the 
Orleans method for wine vinegar. 

11. Bacterium curvum Henneberg. 
(Die deutsche Essigind., No. 11-18, 1906; 
also Cent. f. Bakt., II Abt., 17, 1906, 
791.) From Latin, curvus, bent. 

Rods: 0.4 to 0.5 by 2.0 to 2.4 microns, 



occurring singly or in pairs, cells usually 
oval or elongated, not infrequently sickle- 
shaped, with rounded or pointed ends. 
Not stained with iodine solution. Non- 
motile. 

Wort gelatin : Transparent, round colo- 
nies with raised center and edge, fre- 
quently whitish and dry. 

A non-coherent scanty pellicle is 
formed on the surface of liquid media 
which sinks readily and the liquid is 
quite turbid. 

Forms round white islands on the 
surface of wort with 3 per cent alcohol. 

In old cultures on beer are to be found 
numerous smooth light brown raised 
colonies about 1 mm in diameter on the 
uniform transparent base of the surface 
membrane. 

Acid from arabinose, glucose, raffinose, 
dextrin, ethyl alcohol, propyl alcohol, 
glycerol and erythritol. Small amount 
of acid from fructose, galactose and 
mannitol. No acid from sucrose, maltose 
and lactose (Henneberg, Handbuch d. 
Garungsbakt., 2, 1925, 239). 

Temperature relations : Optimum 25° 
to 30°C. Scant growth at 16° to 17°C. 
No growth at 7° to 8°C. Growth at 35°C. 
No growth at 39°C. 

Source : Isolated from vinegar in the 
quick vinegar process. 

Habitat : Produces acetic acid in the 
quick vinegar process. 

12. Acetobacter viscosum Shimwell. 
{Bacterium aceti viscosum Day and 
Baker, Cent. f. Bakt., II Abt., 36, 1913, 
433; Bacillus aceti viscosum Day and 
Baker, ibid., 437; Also see Baker, Day 
and Hulton, Jour. Inst. Brewing, — , 
1912, 651; Shimwell, Jour. Inst. Brewing, 
42 (N. S. 32), 1936, 586.) From Latin, 
viscous or slimy. 

Rods: 0.4 by 1.2 microns which produce 
ropiness in beer. No capsules observed. 
Non-motile as a rule. Weakly Gram- 
positive. 

Source: From ropy beer. 



FAMILY PSEUDOMONADACEAE 



189 



13. Acetobacter capsulatum Shimwell. 
(Jour. Inst. Brewing, 4^ (N. S. 32), 
1936, 585.) From Latin, capsulated. 

Coccoid rods, 0.8 to 1.0 micron in malt 
extract media. 0.6 to 1.5 microns in other 
media. Produce ropiness in beer. Cap- 
sulated. Motile. Gram-negative. 

Source: From ropy beer. 

14. Acetobacter gluconicum (Her- 
mann).* (Bacterium gluconicum Her- 
mann, Biochem. Zeit., 192, 1928, 198; 
also see Hermann, Biochem. Zeit., 205, 
1929, 297 and Hermann and Neuschul, 
Biochem. Zeit., 233, 1931, 129.) 

It is unfortunate that an organism so 
well described must be placed with other 
species of uncertain standing. How- 
ever, this organism is so closely related 
to the other organisms described in the 
literature that further study is nec- 
essary. 

Source: From kombucha, a mixture of 
fungi and bacteria from tea infusions. 



15. Acetobacter 
Tosic and Walker. 
48, 1942, 82.) 



turbidans Cosbie, 
(Jour. Inst. Brewing, 



This beer vinegar bacterium is char- 
acterized by the production of intense 
turbidity in beer and ale. The descrip- 
tion given does not, at present, warrant 
recognition of the organism as a new 
species. 

Source: From beer. 

16. Bacterium dihydroxyacetonicum 
Virtanen and Biirlund. (Biochem. Zeit., 
169, 1926, 170.) 

There is no adequate description of 
this bacterium, and it is doubtful 
whether it can be properly evaluated 
since various species of Acetobacter also 
possess the ability to produce dihydroxy- 
acetone from glycerol. Consideration 
of this as a nomen nudem was indicated 
by Virtanen to ^'aughn in a personal 
communication in 1938. 

Source: From beet juice. 

17. Acetobacter peroxydans Visser 't 
Hooft. (Inaug. Diss., Delft, 1925, 98.) 

The exact taxonomic position of this 
bacterium will not be clear until further 
comparative studies have been made. 

Source: From hj'drogen peroxide solu- 
tions. 



Genus V. Protamiaobacter den Dooren de Jong.] 

(Bijdrage tot de kennis van het mineralisatieproces. Thesis, Rotterdam, 1926, 
159.) From AI. L., protamine and Latin, bactrum, rod. 

Cells motile or non-motile. Capable of dissimilating alkylamins. Pigmentation 
frequent. Soil or water forms. 

The type species is Protamitiobacter alboflavum den Dooren de Jong. 

Key to the species of genus Protaminobacter. 

I. Non-motile. Gelatin colonies light yellow to colorless. 

1. Protaminobacter alboflavum. 
II. Motile. Gelatin colonies red. 

2. Protaminobacter rubrum. 

1. Protaminobacter alboflavum den Abt., 71, 1927, 218.) From Latin albus, 
Dooren de .Jong. (Thesis, Rotterdam, white; ^at^i/s, yellow. 
1926, 159; also see Cent. f. Bakt., II Rods: Non-motile. Gram-negative. 

* It is uncertain at present who first used this combination. 

t Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, June, 1929; further 
revision by Prof. Robert S. Breed, New York State Experiment Station, Geneva 
New York, April, 1943. 



190 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Gelatin colonies: Circular, dry, light 
yellow or colorless. 

Gelatin stab: No liquefaction. 

Agar colonies: Circular, opaque, pig- 
ment bright red, yellow, light gray or 
colorless. 

Amine agar colonies : Circular, white to 
dark yellow. 

See table below for list of organic sub- 
stances utilized. 



2. ProtaminobacterrubrumdenDooren 
de Jong. (Thesis, Rotterdam, 1926, 
159; also see Cent. f. Bakt., II Abt., 
71, 1927, 218.) From Latin, ruber, red. 

Rods : Motile with single polar flagel- 
lum (Weaver, Samuels and Sherago, 
Jour. Bact., S5, 1938, 59). Gram-nega- 
tive. 

Gelatin colonies : Circular, red, dry. 

Gelatin stab : No liquefaction. 



Table I. — Organic Substances Utilized as a Source of Carbon by Varieties of 
Protaminobacter alboflavum 



ORGANIC ACIDS 

Acetic 

Valerianic 

a-crotonic 

Undecyclic 

Lactic 

|8-oxybutyric 

Succinic 

Formic 

Glutaric 

Adipic 

Fumaric 

Malic 

Tartaric 

Citric 

/S-phenylpropi- 

onic 

Quinic 



+ 

+ + 



+ 


+ 





+ 





+ 


+ 


+ 


+ 


-t- 


-t- 


+ 





+ 





+ 


-1- 


+ 


+ 


+ 








+ 


-!- 



AMINO COxMPOUNDS a 



a-alanin 

a - aminocapronic 

acid 

Leucin 

Propionamid . . 
Capronamid. . . 

Uric acid 

Hippuric acid 

ALCOHOL 

Ethyl 

SUGAR 

Glucose 



a 





7 


a 








+ 


4- 


+ 





+ 





+ 


+ 








+ 





+ 





+ 





+ 





+ 











+ 











+ 


+ 


-h 


+ 


+ 


-t- 


+ 






Ethyl 

Diethyl 

Propyl 

Isopropyl.. . 
Dipropj'l. . . 
Tripropyl... 

Butyl 

Isobutyl 

Diisobutyl. 

Amyl 

Diamyl 

Ethanol 

Glucosamin 
Benzyl 



a 


& 


-y 


+ 


+ 


+ 


-h 


+ 





-l- 


+ 


+ 





+ 





+ 


+ 





+ 








+ 





+ 


+ 


+ 


4- 


+ 


+ 





+ 


+ 


+ 





+ 





+ 


+ 


-f 


+ 


+ 


+ 


+ 





+ 



Catalase is formed. 
Aerobic, facultative. 
Optimum temperature 30 °C. 
Habitat : Soil and water. 

Note : The author recognizes four 
varieties of this species which he dif- 
ferentiates on the basis of organic sub- 
stances attacked (see Table) and pig- 
ment produced. Variety a shows light 
yellow growth on gelatin, bright red on 
agar and yellow on amine agar. Variety ^ 
is light yellow on gelatin, yellow on agar 
and dark yellow on amine agar. Variety 

7 is light yellow on gelatin, light gray on 
agar and yellow on amine agar. Variety 

8 is colorless on gelatin and agar and white 
on amine agar. 



Agar colonies: Circular, red, opaque. 

Amine agar colonies : Circular, dark 
red. 

The following organic acids are at- 
tacked : Acetic, lactic, jS-oxybutyric, 
glycerinic, succinic, malonic, formic, 
methyl formic, glutaric, maleinic, fu- 
maric, malic, tartaric, citric and quinic. 

The following amino compounds are 
attacked: Sarcosin, betain, hippuric 
acid, asparagine, propionamid, caprona- 
mid, lactamid, succinamid, allantoin 
and uric acid. 

Glucose is fermented. 

Catalase is formed. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Habitat : Soil and water. 



FAMILY PSEUDOMONADACEAE 



191 



Genns VI. Mycoplana Gray and Thornton * 

(Cent. f. Bakt., II Abt., 73, 192S, S2.) From Greek, mykes, fungus; plane, a wan- 
derer or traveller. 

Cells branching, especially in 3'oung cultures. Frequently banded when stained. 
Capable of using phenol and similar aromatic compounds as a sole source of energy. 
Grow well on standard culture media. 

Type species Mycoplana dimorpha Graj^ and Thornton. 

Key to the species of genus Mycoplana. 

I. Gelatin not liquefied. 

1. Mycoplana dimorpha. 
II. Gelatin liquefied. 

2. Mycoplana hxdlata. 



1. Mycoplana dimorpha Gray and 
Thornton. (Cent. f. Bakt., II Abt., 
73, 1928, 82.) From Greek, di, two; 
morphos, forms. 

■ Short, curved and irregular rods, 0.5 
to 0.7 by 1.25 to 4.5 microns, showing 
branching especially in young cultures. 
fMotile, with long polar flagella. Gram- 
negative. 

Gelatin colonies : Circular, buft', smooth, 
resinous, entire. 

Gelatin stab : Xo liquefaction. Growth 
filiform. 

Agar colonies: Circular, buff, convex, 
smooth, glistening, entire. 

Agar slant: Filiform, white, convex, 
glistening, entire. 

Broth: Turbid, with surface ring. 

Nitrites not produced from nitrates 
but gas evolved in fermentation tubes. 

Starch hydrolyzed. 

No acid from carbohydrate media. 

Attacks phenol. 

Aerobic. 

Optimum temperature below 30 °C. 

Source: One strain isolated from soil. 

Habitat: Probably in soil. 



2. Mycoplana bullata Gray and Thorn- 
ton. (Cent. f. Bakt., II Abt., 73, 1928, 
83.) From Latin, bullatus, furnished 
with a boss or knob. 

Rods, curved and irregular, branching, 
0.8 to 1.0 by 2.25 to 4.5 microns. fMotile 
with polar flagella. Gram-negative. 

Gelatin colonies : Circular, buff, smooth, 
glistening, edge diffuse. Partially lique- 
fied. 

Gelatin stab: Saccate liquefaction. 

Agar colonies : Circular, white, convex, 
smooth, glistening, entire. 

Agar slant: Filiform, white, convex, 
smooth, glistening, entire. 

Broth: Turbid. 

Nitrites not produced from nitrates. 
Gas, presumabl}^ N, in fermentation 
tubes. 

Starch not hydrolyzed. 

No acid from carbohj'drate media. 

Attacks phenol. 

Aerobic. 

Optimum temperature below 30 °C. 

Source: Two strains isolated from soil. 

Habitat: Probably in soil. 



* Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, June, 1929. 

t The original statements regarding the flagellation of the.se species are contra- 
dictory. The first reads "Polar, peritrichous; the second "Polar or peritrichous". 
Drawings given usually indicate peritrichous rather than polar flagellation. Further 
study is needed before these species can be properly placed in relation to other known 
species. — Editors. 



192 MANUAL OF DETERMINATIVE BACTERIOLOGY 

TRIBE II. SPIRILLEAE KLUYVER AND VAN NIEL. 

(Cent. f. Bakt., II Abt., H, 1936, 346.) 
More or less spirally curved cells. 

Key to the genera of tribe Spirilleae. 

I. Generally motile by means of a single polar flagellum. 

A. Short, bent rods occurring singly or united into spirals. 

Genus I. Vibrio, p. 192. 

B. Slightly curved rods of variable length. Strict anaerobes which reduce 

sulfates to hydrogen sulfide. 

Genus II. Desulfovibrio , p. 207. 

C. Cells oxidize cellulose forming oxj^cellulose. Growth on ordinary culture 

media is feeble. 

1. Long, slightly curved rods with rounded ends. 

Genus III. Cellvibrio, p. 209. 

2. Short, curved rods with pointed ends. 

Genus IV. Cellfalcicula, p. 211. 
II. Generally motile by means of a tuft of polar flagella. Cells of varying thick- 
ness, and length and pitch of spiral, forming either long curves or portions 
of a turn. 

A. Oxidize inorganic sulfur compounds. Cells contain free sulfur granules. 

Genus V. Thiospira, p. 212. 

B. Not as above. 

Genus VI. Spirillum, p. 212. 

Genus I. Vibrio Muller* 

(Miiller, Vermium terrestrium et fluviatilum, 1, 1773, 39; Pacinia Trevisan, Att^ 
d. Accad. Fisio-Medico-Statisticain Milano, Ser. 4, 3, 1885, 83; Microspira Schroeter, 
in Cohn, Kryptogamen-Flora von Schlesien, 3, 1, 1886, 168; Pseudospira De Toni and 
Trevisan, Sylloge Fungorum, 8, 1889, 1018; Photobacterium Beijerinck, Arch, neerl. 
d. sci. exactes, 23, 1889, 401; Liquidovibrio Orla-Jensen, Cent. f. Bakt. II Abt., 22, 
1909, 333; Solidovibrio Orla-Jensen, ibid.; Dicrospira Enderlein, Sitzber. Ges. naturf. 
Freunde, Berlin, 1917, 313.) From Latin, vibro, vibrate. 

Cells short, curved, single or united into spirals. Motile by means of a single polar 
flagellum which is usually relatively short; rarely, two or three flagella in one tuft. 
They grow well and rapidly on the surface of standard culture media. Aerobic to 
anaerobic species. Mostly water forms, a few parasites. 

The type species is Vibrio comma (Schroeter) Winslow et al. 

Key to the species of genus Vibrio. 

I. Gelatin liquefied. 

A. Nitrites produced from nitrates. 
1. Indole is formed. 

a. Milk not coagulated. 

1. Vibrio comma. 

2. Vibrio berolinensis. 
aa. Milk coagulated. 

3. Vibrio metschnikovii. 

* Revised by Prof. D. H. Bergey, Philadelphia, Pennsylvania, April, 1937; partial 
revision by Capt. Wm. C. Haynes, Sn.C, Fort Bliss, Texas, July, 1943 and by Lt. 
Col. A. Parker Kitchens, University of Pennsylvania, Philadelphia, Penna., Decem- 
ber, 1943. 



FAMILY PSEUDOMONADACEAE 



193 




2. Indole not formed. 

a. Milk not coagulated. 

4. Vibrio tyrogenus. 

5. Vibrio xenopus. 
B. Nitrites not produced from nitrates. 

1. Indole is formed. 

a. Milk coagulated, peptonized. 

6. Vibrio piscium. 

2. Indole not formed. 

a. Milk acid, coagulated. 

7. Vibrio proteus. 

8. Vibrio wolfii. 

9. Vibrio sputigeniis. 

10. Vibrio liquefaciens. 
aa. Milk not coagulated. 

b. Growth on potato thin, barely visible. 

11. Vibrio strictus. 
bb. Xo growth on potato. 

12. Vibrio aquatilis. 
aaa. Action on milk not reported. 

b. Acid from glucose. Attacks naphthalene. 

13. Vibrio neocistes. 

bb. No acid from carbohydrates. Attacks naphthalene. 

14. Vibrio cuneatus. 

bbb. No acid from carbohydrates. Liquefies agar. 

15. Vibrio granii. 
II. Gelatin not liquefied. 

A. Nitrites produced from nitrates. 

1. Acid and gas from glucose. 

16. Vibrio leonardii. 

2. Acid but not gas from glucose. Liquefies agar. 

17. Vibrio agarliquefaciens. 

B. Nitrites not produced from nitrates. 

1. Acid from glucose. 

IS. Vibrio cyclosiles. 

2. No acid from carbohydrates. 

19. Vibrio percolans. 

C. Nitrite production not reported. 

1. Requires the addition of ammonium sulfate for growth. Ammonium 

sulfate agar liquefied. 

20. Vibrio andoi. 

2. Do not require ammonium sulfate for growth. 

a. Indole not formed. 

b. Microaerophilic, becoming aerobic. 

21. Vibrio fetus. 
bb. Aerobic, facultative. 

22. Vibrio pierantonii . 



1. Vibrio comma (Schroetcr) Wins- 479; Spirillum cholerae asiaticae Zopf, 
low et al. (Kommabacillus, Koch, Die Spaltpilze, 3 Aufl., 1S85, 69; Paci/i/a 
Berliner klin. Wochenschr., 21, 1884, c/ioieraeas/aiicae Trevisan, Atti d. Accad. 



194 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Fisio-Med.-Statistica in Milano, Ser. 4, 3, 
1885, 84; Microspira comma Schroeter, in 
Cohn, Kryptogamen Flora v. Schlesien, 
3, 1, 1886, 168; Vibrio cholerae Neisser, 
Arch. f. Hyg., 19, 1893, 199; Vibrio 
cholerae asiaiicae Pfeiffer, in Fliigge, Die 
Mikroorganismen, 2, 1896, 527; Winslow 
et al., Jour. Bact., 5, 1920, 204; Bacillus 
cholerae Holland, Jour. Bact., 5, 1920, 
217; Bacillus comma Holland, ibid., 218; 
Spirillum cholerae-asiaticae Holland, 
ibid., 225; Vibrio cholerae-asiaticae Hol- 
land, ibid., 226.) From Latin, comma. 

Slightly curved rods, 0.3 to 0.6 by 1.0 
to 5.0 microns, occurring singly and in 
spiral chains. Cells may be long, thin 
and delicate or short and thick. May 
lose their curved form on artificial culti- 
vation. Motile, possessing a single 
polar flagellum. Gram-negative. 

Gelatin colonies: Smajl, yellowish- 
white. 

Gelatin stab : Rapid napiform lique- 
faction. 

Agar colonies : Circular, whitish-brown, 
moist, glistening, translucent, slightly 
raised, entire. 

Agar slant: Brownish-gray, moist, 
glistening. 

McConkey's medium: Good growth, 
colonies colorless when young, soon 
pinkish, medium becomes darker red. 

Broth: Slightly turbid, with fragile, 
wrinkled pellicle and flocculent precipi- 
tate. 

Peptone water: Characteristic rapid 
growth, chiefly at surface, where after 
6 to 9 hours, a delicate membrane is 
formed; little turbidity, deposit appar- 
ently derived from pellicle (Topley and 
Wilson, Princip. Bact. and Immun., 
2nd ed., 1936, 388). Readily isolated 
from the surface film of 0.1 per cent 
peptone water. 

Litmus milk : Alkaline at the top and 
slightly acid at bottom; generally not 
coagulated; peptonized; reduced. 



Potato : Dirty -white to yellowish, moist, 
glistening, spreading. 

Blood serum: Abundant growth, some- 
times slow liquefaction. 

Blood agar: The blood pigment is di- 
gested forming a greenish zone around 
colonies; a true soluble hemolysin is 
not formed (the El Tor vibrio also di- 
gests blood pigment but in addition pro- 
duces a soluble hemolysin. Otherwise it 
is said to be indistinguishable from the 
typical cholera vibrio) . 

Indole is formed. 

Nitrites produced from nitrates. 

Cholera-red reaction, which depends 
on production of indole and reduction of 
nitrates is positive. 

Hydrogen sulfide is formed. 

Acid but not gas from glucose, fruc- 
tose, galactose, maltose, sucrose and 
mannitol. Slowly from glycerol. Does 
not attack lactose, inulin or dulcitol. 

Group I of Heiberg (Classification of 
Vibrio cholerae and Cholera-like Vibrios. 
Copenhagen, 1935) ferments mannose 
and sucrose but not arabinose. 

Hydrolyzes starch actively in alkaline 
media. 

High alkali but low acid tolerance; 
optimum pH 7.6 to 8.0; for isolation on 
Dieudonne's medium pH 9.0 to 9.6. 

Aerobic, grows best in abundant oxy- 
gen; under strict anaerobiosis may fail 
to grow altogether. 

Optimum temperature 37 °C. Maxi- 
mum 42°C. Minimum 14°C. 

Source: From intestinal contents of 
cholera patients in Egypt and India. 

Habitat : Intestinal contents of cholera 
patients and carriers. 

The relationships existing among the 
cholerigenic and non-pathogenic water 
vibrios, although studied intensively, 
have not yet been completely defined. 
As a working scheme, based on somatic 
(O) and flagellar (H) antigen studies, 
Gardner and Vankatraman (Jour. Hyg., 
35, 1935, 262-282) suggest the following 



FAMILY PSEUDOMONADACEAE 



195 



Cholera group of vibrios. 
(Biochemically similar. Common H antigen.) 



0-sub-group I. 



Non-hemolytic 
(goat cells). 
Cholera vibrios. 
Types — original, 
variant and 
middle. 



Hemolytic (goat 

cells). 

El Tor vibrios. 

Types — original 

and variant 

(Pmiddle). 



Linton (Bact. Rev., 4, 1940, 275) has 
outlined a classification of the vibrios 
based upon their protein and poly- 
saccharide structures. Using chemical 
methods, it was found that one poly- 
saccharide and one protein was com- 
monly obtained from each strain of 
vibrio; when exceptions occurred, it was 
invariably noted that the strain was 
undergoing dissociation. Given a single 
protein and polj'saccharide in each 
vibrio, it was possible to divide the 
strains into six groups, which were 
numbered in the order of their discovery 
as shown in the table. 

A chemical grouping of the cholerigcnic 
and water vibrios. 



Group 


Protein Type 


I 


I 


II 


I 


III 


II 


IV 


II 


V 


II 


VI 


I 



Polysaccharide 
Type 



I 

II 

II 

I 

III 

III 



The strains of Groups I and II possess 
the same protein and different poly- 
saccharides. These are derived from 
cases of cholera and have the serological 



O sub-groups II, HI, IV, V, VI and 

individual races (mostly hemolytic). 
Paracholera, cholera-like, and some 
El Tor vibrios. 



(Types \Yithin sub-groups underlined.) 



and biochemical characteristics of O- 
Group I, Vibrio cholera. Group I strains 
are more common than those of Group 
II, which have, however, been isolated 
from epidemics with a high mortality. 
The phospholipid fraction is common to 
both types when isolated in the early 
part of an epidemic, but is not found in 
strains of other groups. The harmless 
water vibrios, which are so heteroge- 
neous serologically (Taylor and Ahuja, 
Indian Jour. Med. Res., 26, 1938, 8-32) 
form a single chemical group with a 
homogeneous structure. They fall into 
Group III, which differs in its protein 
structure from the authentic cholera 
vibrios, and resembles Group II in its 
polysaccharide. The vibrios of Group 
IV, which came from El Tor and from 
chronic vibrio carriers are believed on 
epidemiological grounds to be harmless, 
although serological methods have failed 
to distinguish them from cholerigcnic 
vibrios. Group V, which, like III and 
IV, contains protein II, consists, like 
Group IV, of strains from chronic vibrio 
carriers. Group VI strains are only 
rarely isolated in nature and representa- 
tives of this group are generally found 
among collections of old laboratory 
strains. They appear to be the result of 
polysaccharide variation from Group I 



196 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



after long-continued growth on artificial 
media. 

2. Vibrio berolinensis Neisser. (Arch, 
f. Hyg., 19, 1S93, 200; Microspira bero- 
linensis Migula, in Engler and Prantl, 
Die natiirl. Pflanzenfam., 1, la, 1895, 33.) 
From M. L., the genitive of Berolina, the 
Latin name for Berlin. 

Curved rods, somewhat smaller than 
Vibrio comma. Frequently occurring 
in pairs. Motile, possessing a polar 
flagellum. Pleomorphic. Gram-nega- 
tive. 

Gelatin colonies: Small, grayish, 
slightly granular, fragmented. 

Gelatin stab: Slow, napiform lique- 
faction. 

Agar slant: Grayish-yellow, moist, 
glistening. 

Broth: Turbid, with gray pellicle. 

Litmus milk: No coagulation, no acid. 

Potato: Brownish streak. 

Indole is formed. 

Nitrites produced from nitrates. 

Not pathogenic for mice, pigeons or 
guinea pigs. 

Aerobic, facultative. 

Optimum temperature 37 °C. Mini- 
mum above 10 °C. Maximum less than 
60 °C. 

Source : Isolated from filtered Spree 
river water. 

3. Vibrio metschnikovii Gamaleia. 

(Gamaleia, Ann. Inst. Pasteur, 2, 1888, 
482; Pacinia meischmkoffi. Trevisan, I 
generi e le specie delle Battcriacee, 1889, 
23; Spirillum metschnikovi Sternberg, 
Man. of Bact., 1893, 511; Vibrio nord- 
hafen Pfuhl, Ztschr. f. Hyg., 22, 1894, 
234; Microspira metschnikoffii Migula, 
in Engler and Prantl, Die nattirl. Pfian- 
zenfam., 1, la, 1895, 33.) Named for 
Metschnikoff, Russian bacteriologist. 

Probable synonyms : Vibrio schuyl- 
kiliensis Abbott, Jour. Exp. Med., 1, 
1896, 424 {Microspira schxylkiliensis 
Chester, Manual Determ. Bact., 1901, 
334); Vibrio dannbicus Heidcr, Cent. f. 



Bakt., H, 1893, 341 {Microspira danubica 
Migula in Engler and Prantl, Die natiirl. 
Pflanzenfam., 1, la, 1895, 33; Spirillum 
danubicum Holland, Jour. Bact., 5, 1920, 
225). 

Curved rods, somewhat shorter and 
thicker than Vibrio comma. Motile. 
Gram-negative. 

Gelatin colonies : Like those of Vibrio 
comma. 

Gelatin stab: Rapid, napiform lique- 
faction. 

Agar slant: Yellowish, plumose, moist, 
glistening. 

Broth: Turbid, with thin, white pel- 
licle. 

Litmus milk: Acid, coagulated (eighth 
day); not peptonized. 

Potato: Delicate, brownish growth. 

Indole is formed. 

Nitrites produced from nitrates. 

Pathogenic for pigeons, fowls, and 
guinea pigs. 

Aerobic, facultative. 

Optimum temperature 37°C. Maxi- 
mum less than 45°C. 

Source : Isolated from fowl dead of a 
choleraic disease. 

Habitat: The intestinal contents of 
chickens, pigeons and other animals suf- 
fering from a cholera-like disease. 

4. Vibrio tyrogenus (Fliigge) Hol- 
land. (Kasespirillen, Deneke, Deutsch. 
med. Wochnschr., 11, 1885, 33; Spirillum 
tyrogenum Fliigge, Die Mikroorganismen, 
2 Aufl.. 1886, 386; Pacinia denekei Trevi- 
san, I generi e le specie delle Batteriacee, 
1889, 23; Microspira tyrogena Migula, 
in Engler and Prantl, Die natiirl. Pflan- 
zenfam., 1, la, 1895, 33; Holland, Jour. 
Bact., 5, 1920, 225; Vibrio denekei 
Ilauduroy et al., Diet. d. Bact. Path., 
1937, 541.) From Greek tyros, cheese; 
genes, produced from. 

Curved rods, rather smaller and more 
slender than Vibrio comma, often very 
long, closely wound spirals. Motile, 
possessing a polar flagellum. Gram- 
negative. 



FAMILY PSEUDOMONADACEAE 



197 



Gelatin colonies : Small, gray, granular, 
entire. 

Gelatin stab: Rapid, saccate liquefac- 
tion. 

Agar slant: Yellowish-white, plumose, 
glistening. 

Broth: Turbid. 

Litmus milk : Not coagulated. 

Potato: No growth. 

Indole not formed. 

Slight production of nitrites from 
nitrates. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Source: Isolated from cheese. 

5. Vibrio xenopus Schrire and Green- 
field. (Trans. RoA^al Soc. So. Africa, 
17, 1930, 309.) From Xenopus, a genus 
of African toads. 

Spiral forms, occurring singly and in 
pairs. Non-motile. Gram-negative. 

Gelatin stab : Slow, cratcriform liciue- 
faction. 

Agar colonies : Small, white, glistening, 
slim}% entire. 

Agar slant : Grayish-white, slimy, 
entire. 

Broth: Turbid with flocculent sedi- 
ment. 

Litmus milk: Unchanged. 

Potato: Not reported. 

Indole is not formed. 

Nitrites are produced slowly from 
nitrates. 

Blood serum is peptonized. 

Starch is not hydrolj'zed. 

Acid from glucose, fructose, maltose, 
glycerol and sorbitol. 

Aerobic, facultative. 

Optimum temperature 37°C. 

Source : Found in abscess of pectoral 
muscle of African toad. 

6. Vibrio piscium David. (Cent. f. 
Bakt., I Abt., Orig., 102, 1927, 46.) From 
Latin pise is, fish. 

Curved rods : 0.3 to 0.5 by 2.0 microns. 
Motile with a single polar flagellum. 
Gram-negative. 



Gelatin colonies : Circular, granular, 
opaque. 

Gelatin stab : Napiform liquefaction. 

Agar colonies : Yellowish, circular, 
smooth, entire, iridescent. 

Agar slant : Light yellow, transparent 
streak. 

Broth: Slight turbidity, with thin 
pellicle. 

Litmus milk : Soft coagulum. Pep- 
tonized, alkaline. 

Potato: Brownish-red streak. 

Indole is formed. 

Nitrites not produced from nitrates. 

Hydrogen sulfide formed. 

No action in sugar media. 

Pathogenic for frogs. 

Aerobic, facultative. 

Optimum temperature 18° to 20°C. 

Habitat : Causes epidemic infection in 
fish. 

7. Vibrio proteus Buchner. (Kom- 
mabacillus der cholera nostras, Finkler 
and Prior, Deutsche med. Wochenschr., 
1884, 632; Buchner, Sitzungsber. d. 
Gesel. f. Morph. u. Physiol., Miinchen, 
Heft 1, 1885, 10; Pacinia finkleri Trev- 
isan, Atti d. Accad. Fi sio -Medico - 
Statistica in Milano, Ser. 4, 3, 1885, 84; 
Microspira finkleri Schroeter, in Cohn, 
Krytogamen-Flora von Schlesien, 3, 1, 
1886, 168; Spirillum finkleri Crookshank, 
Man. of Bact., 3rd ed., 1890, 282; Micro- 
spira protea Chester, Manual of Deter- 
minative Bacteriology, 1901, 338; Vibrio 
finkleri Holland, Jour. Bact., 5, 1920, 
225.) From Greek, Proteus, a marine 
deity who had the power of assuming any 
shape he chose. 

Curved rods : 0.4 to 0.6 by 2.4 microns, 
often pointed at both ends. Motile, 
possessing a polar flagellum. Gram- 
negative. 

Gelatin colonies: Small, gray, circular, 
granular, entire. 

Gelatin stab: Rapid, saccate lique- 
faction. 

Agar slant : Dirty grayish, plumose. 

Broth: Turbid, with fetid odor. 



198 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Litmus milk : Slightly acid ; coagulated ; 
peptonized. 

Potato : Grayish, slimy layer. 

Indole not formed. 

Nitriites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 30 °C. 

Source: Isolated from feces of pa- 
tients suffering from cholera nostras. 

Habitat : Intestinal contents in cholera 
nostras and cholera infantum. 

8. Vibrio wolfii (Migula) Bergey et al. 
{Bacillus choleroides Wolf, Mlinch. med. 
Wochenschr., 40, 1S93, 693; Microspira 
wolfii Migula, Syst. d. Bakt., 2, 1900, 
1001; not Microspira choleroides Migula, 
loc. cit., 992; Bergey et al., Manual, 1st 
ed., 1923, 80.) Named for Wolf, who 
first isolated this organism. 

Curved rods and S-shaped forms. 
Motile. Gram-negative. 

Gelatin colonies: Small, grayish-white, 
spreading. 

Gelatin stab: Infundibuliform lique- 
faction. 

Agar slant: Gray, moist layer. 

Broth: Turbid, witli gray pellicle. 

Litmus milk: Acid; coagulated. 

Potato : Yellowish-white layer. 

Blood serum: Rapid liquefaction. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 37 °C. 

Source : Isolated from cervical secre- 
tions in chronic endometritis. 



Gelatin colonies: Small, circular, 
slightly granular, yellowish, becoming 
brownish. 

Gelatin: Crateriform liquefaction. 

Agar slant: Grayish-white, moist. 

Broth: Turbid, no pellicle formed. 

Litmus milk: Acid; coagulated. 

Potato : Thin, gray layer, spreading. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 37 °C. 

Source : Isolated from sputum. 

10. Vibrio liquefaciens (Migula) Ber- 
gey et al. (Bonhoff, Arch. f. Hyg., 19, 
1893, 248; Microspira liquefaciens Mi- 
gula, Syst. d. Bakt., 2, 1900, 990; Bergey 
et al.. Manual, 1st ed., 1923, 81.) From 
Latin, liquefacio, to make liquid. 

Comma and S-shaped rods. Motile. 
Gram-negative. 

Gelatin colonies: Circular, with ir- 
regular margin, surrounded by a rose- 
colored zone. 

Gelatin stab : Slow, napiform lique- 
faction. 

Agar slant: Smooth, grayish, plumose. 

Broth : Turbid, with heavy grayish 
pellicle. 

Litmus milk: Acid; coagulated. 

Potato : Moist, brownish layer. 

Indole is not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 37 °C. 

Habitat : Water. 



9. Vibrio sputigenus (Migula) Bergey 
et al. (Vibrio aus Sputum, Brix, Hyg. 
Rundschau, 4, 1894, 913; Microspira 
sputigena Mignla, Syst. d. Bakt., f, 1900, 
981; Bergey ct al.. Manual, 1st ed.. 1923, 
80.) From Latin, spuo {sputus), spu- 
tum; -genes, produced from. 

Slightly curved rods, about the same 
size and form as Vibrio comma, occurring 
singly, occasionally three or four in a 
chain. Motile. Possessing a polar fiagel- 
lum. Gram-negative. 



11. Vibrio strictus Kutscher. (Ztschr. 
f. Hyg., 19, 1895, 469.) From Latin 
stringo (strictus), constricted. 

Markedly curved rods, of about twice 
the size of Vibrio comma. Motile. 
Gram-negative. 

Gelatin colonies: Small, circular, yel- 
lowish, with serrate margin. 

Gelatin stab: Slow, napiform to sac- 
cate liquefaction. 

Agar slant : Growth plumose, moist. 

Broth: Turbid, with gray pellicle. 



FAMILY PSEUDOMONADACEAE 



199 



Litmus milk: Not coagulated. 
Potato: Thin, barely visible layer. 
Blood serum is slowlj' liquefied. 
Indole is not formed. 
Nitrites not produced from nitrates. 
Pathogenic for guinea pigs. 
Aerobic, facultative. 
Optimum temperature 37 °C. 
Habitat : Water. 

12. Vibrio aquatilis Giinther. (Deut- 
sche med. Woehenschr., 1S92, 1124; 
M icrospira aquatilis Migula, System der 
Bakterien, 2, 1900, 993.) From Latin, 
aquaticus, living in water. 

Curved rods, like Vibrio comma. 
Motile, possessing a polar flagellum. 
Gram-negative. 

Gelatin colonies: Circular, brownish, 
finely granular, entire. 

Gelatin stab : Crateriform liquefaction. 

Agar slant : Moist, grayish, glistening. 

Broth: Slightly turbid. 

Litmus milk : Not coagulated. 

Potato: No growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 30 °C. 

Habitat : Water. 

13. Vibrio neocistes Gray and Thorn- 
ton. (Gray and Thornton, Cent. f. 
Bakt., II Abt., 73, 1928, 92.) From 
Greek neos, new and kiste bo.x or ark. 
Here used as the equivalent of Newark, 
the name of a city in England. 

Curved rods: 0.5 to 1.0 by 1.0 to 4.0 
microns. Motile with one to three 
polar flagella. Gram stain not recorded. 

Gelatin colonies : Liquefied. 

Gelatin stab : Liquefied. Medium red- 
dened. 

Agar colonies : Circular or amoeboid, 
buff to brownish, convex, smooth, glis- 
tening, entire. 

Agar slant :Filiform, fluorescent, raised, 
smooth, glistening, undulate. 

Broth: Turbid. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 



Acid from glucose. 
Attacks naphthalene. 
Aerobic, facultative. 
Optimum temperature. 
Habitat: Soil. 

14. Vibrio cuneatus Gray and Thorn- 
ton. (Gray and Thornton, Cent. f. Bakt., 
II Abt., 73, 1928, 92.) From Latin, 
cuneo (cuneatus) wedge. 

Curved rods : 1.0 by 1.0 to 3.0 microns, 
the cells tapering at one extremity. 
Motile with one to five polar flagella. 
Gram-negative. 

Gelatin colonies: Liquefied. 

Gelatin stab: Liquefied. 

Agar colonies : Circular to amoeboid, 
white to buff, fiat to convex, smooth, 
translucent, border entire. 

Agar slant: Filiform, whitish, smooth, 
glistening. 

Indole not recorded. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

No acid from carbohydrate media. 

Attacks naphthalene. 

Aerobic, facultative. 

Optimum temperature 30° to 35°C. 

Source : One strain isolated from 
soil from Rothamsted, England. 

Habitat: Soil. 

15. Vibrio granii (Lundestad) Stanier. 
(Bacterium granii Lundestad, Cent. f. 
Bakt., II Abt., 75, 1928, 330; Achromo- 
bacter granii Bergey et al., Manual, 3rd 
ed., 1930, 222; Stanier, Jour. Bact., 42, 
1941, 538.) Named for Prof. H. H. Gran, 
who first detected agar-liquefying bac- 
teria. 

Rods: 0.6 to 0.8 by 1.4 to 2.4 microns, 
with rounded ends, occurring singly, in 
pairs, and at times in short chains. 
Motile. Polar flagellate (Stanier, loc. 
cit.). Gram-negative. 

Fish-gelatin colonies : Punctiform, 
black, glistening. 

Fish-gelatin stab : Slow, crateriform 
liquefaction. 

Sea-weed agar colonies: Circular, flat 



200 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



opaque, glistening, white, slimy, entire. 
Agar is dissolved. 

Fish-agar slant: Flat, white, elevated, 
glistening, undulate. Liquefied. 

Broth: Turbid with grayish-white, 
slimy sediment. 

Indole not formed. 

Nitrites not produced from nitrates. 

Starch usually hydrolyzed. 

No action on sugars. 

Aerobic, facultative. 

Optimum temperature 20° to 25 °C. 

Source : Sea water of Norwegian Coast. 

Habitat: Presumably sea water and 
on sea weeds. 

16. Vibrio leonardii Metalnikov and 
Chorine. (Ann. d. I'Inst. Pasteur, ^2, 
1928, 1647.) Named for Leonard. 

Curved rods with rounded ends, 0.5 
to 1.0 by 2.0 to 3.0 microns. Motile 
with 1 to 3 polar flagella. Gram-negative. 

Gelatin stab : No liquefaction. 

Agar colonies: Small, transparent, cir- 
cular, having a characteristic odor. 

Broth: Turbid, with thin pellicle. 

Litmus milk: No coagulation, acid, 
with reduction of litmus. 

Potato: Slight, colorless growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Blood serum not liquefied. 

Hydrogen sulfide formed. 

Acid and gas from glucose, fructose, 
galactose, lactose, sucrose and mannitol. 
No acid or gas from maltose or glycerol. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Habitat : Highly pathogenic for insects 
as Galleria mellonella L. (bee moth), 
and Pyrausta nubialis Hiibn. (European 
corn borer). 

17. Vibrio agarliquefaciens (Gray and 
Chalmers) Bergey et al. (Microspira 
agar-liquefaciens Gray and Chalmers, 
Ann. Appl. Biol., 11, 1924, 325; Manual, 
4th ed., 1934, 119.) From Latin, lique- 
facio, liquefying; Malay, agar, a jelly 
from seaweeds. 

Short curved rods, usually c-shaped, 



with occasional s-shaped and coccoid 
forms. Cells 2.0 microns long by 0.5 
to 0.7 micron broad; 3.0 to 5.0 microns 
long in division stages. Coccoid forms 
stained, 0.5 to 0.7 micron long. Motile 
with a single polar flagellum. Gram stain 
not reported. 

Gelatin stab : Very slight surface 
growth after one month ; the streak then 
shows a beaded line. No liquefaction. 

Agar colonies : Surface colonies appear 
as a whitish growth in a depression, sur- 
rounded by a white ring. The colony is 
later surrounded by a ring of liquid agar. 
Deep colonies show a clear area and may 
be irregular, oval or angular. 

Agar slant : A deep groove is cut along 
the inoculation streak, whitish growth 
along sides. The gel is later much weak- 
ened. 

Broth: Slightly turbid. No pellicle. 

Acid from glucose, lactose and mal- 
tose. No acid from sucrose or glycerine. 

Utilizes ammonia salts as a source of 
nitrogen. 

Decomposes cellulose and agar. The 
presence of one per cent glucose pre- 
vents the liquefaction of agar. 

Nitrites produced from nitrates. 

Starch is hydrolyzed. 

Aerobic. 

Temperature relations: Optimum 25° 
C, will grow at 16° but not at 34 °C. 

Habitat: Soil. 

18. Vibrio cyclosites Gray and Thorn- 
ton. (Gray and Thornton, Cent, fiir 
Bakt., II Abt., 73, 1928, 92.) From 
Greek kyklos, circle or ring; sited, to eat; 
M. L. cyclosites, feeding on rings, i.e. ring 
compounds. 

Curved rods: 0.5 to 1.0 by 1.5 to 4.0 
microns. Motile with a single polar 
flagellum. Gram-negative. 

Gelatin colonies : Circular, buff to 
brown, flat, smooth, glistening, entire. 

Gelatin stab: No liquefaction. 

Agar colonies : Circular to irregular, 
pale buff (later greenish), smooth, entire. 

Agar stab: Filiform, greenish buff, 
raised, smooth, undulate. 



FAMILY PSEUDOMOXADACEAE 



201 



Broth : Turbid. 

Indole not reported. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Acid from glucose. 

Attacks phenol and ?n-cresol. 

Aerobic, facultative. 

Optimum temperature 30° to 35 °C. 

Habitat : Soil. 

19. Vibrio percolans Mudd and War- 
ren. (Jour, of Bact., 8, 1923, 447.) 
From Latin, percolo (percolatus) , filter- 
ing. 

Curved rods: 0.3 to 0.4 by 1.5 to 1.8 
microns, occurring singly or in short 
chains. Pleomorphic. Actively motile 
by means of 1 to 3 polar flagella. Gram- 
negative. 

Gelatin stab: Xo liquefaction. 

Agar colonies : Circular, slightly con- 
vex, amorphous, entire. 

Agar slant : Bluish-white, glistening, 
streak. 

Broth: Turbid. Pellicle, sediment. 

Litmus milk : L'nchanged. 

Potato: White, slimy streak. 

Indole not formed. 

Nitrites not produced from nitrates. 

Blood serum not licjucfied. 

Starch not hydrolj'zed. 

No action on carbohydrates. 

Passes through bacterial filters. 

Aerobic, facultative. 

Optimum temperature 30°C. 

Non-pathogenic. 

Source: Isolated from haj^ infusion. 

Habitat: Water. 

20. Vibrio andoi Aoi and Orikura. 
(Cent. f. Bakt., II Abt., 74, 1928, 331.) 
Named for Andoi, a Japanese scientist. 

Curved rods, with more or less tapering 
ends, c- or s-shaped, 0.5 to 0.8 by 1.5 to 
2.5 microns. Motile, with a single polar 
flagellum. Gram-negative. 

Gelatin: No growth. 

Agar media : No growth. 

Broth: No growth. 

Litmus milk : No growth. 

Potato : No growth. 



Ammonium sulfate agar colonies : 
Punctiform, circular, concave, surrounded 
with clear zone. 

Ammonium sulfate agar slant : Grayish, 
becoming straw-yellow, sinking into the 
medium as the agar liquefies. 

Cellulose media: No growth. 

Starch hydrolyzed. 

Glucose, fructose, galactose, mannose, 
xylose and "honyak" are fermented. 

Xylan is decomposed. 

Cellobiose is decomposed. 

Aerobic, facultative. 

Optimum temperature 25 °C. Mini- 
mum 8 °C . Maximum 37 °C . 

Source : Rotted stable manure. 

Habitat: Presumably decomposing or- 
ganic matter. 

21. Vibrio fetus Smith and Taylor. 
(Spirillum causing abortion in sheep, 
^lacFadyean and Stockman, Rept. Dept. 
Comm. Ministr\' Agric. on Epizootic 
Abortion, London, 1909, 156; ibid., 1913, 
111 ; Spirillum associated with infectious 
abortion, Smith, Jour. Exp. Med., 28, 

1918, 701; Smith and Taylor, ihid., 30, 

1919, 299; Spirillum fetus Lehmann and 
Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 
552; Vibrio foetus ovis Buxton, First 
Rept. of Director, L^niv. Cambridge 
Instit. Animal Pathol., 1929-1930, 47.) 
From li. foetus, fetus. 

Curved rods : The smallest forms ap- 
pear as minute curved s-shaped lines, 
other forms very long; 0.2 to 0.5 by 1.5 
to 5.0 microns. Motile by means of one, 
rarely two, polar flagella. Occasionally 
forms capsules. Granules present in 
older cultures. Gram-negative. 

Gelatin: Xo growth. 

Agar slant: No surface growth by 
freshly isolated strains. Laboratory 
strains produce a scanty, grayish-white, 
glistening surface growth. 

Subsurface agar colonies : Small, yel- 
low, opaque. 

Broth : A viscid ring pellicle may ap- 
pear, faint clouding of the medium occurs ; 
a filmy, stringy deposit may settle out. 

Litmus milk: No growth 



202 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Potato: No growth. 

Indole not formed. 

Nitrite production not reported. 

Blood serum slant: Feeble growth. 
No liquefaction. 

No gas from carbohydrates. No change 
or slightly acid from glucose, lactose 
and sucrose. 

Optimum temperature 37°C. With- 
stands 55°C for 5 minutes. 

Aerobic or microaerophilic. 

Pathogenesis : Causes abortion in cattle. 

Source : Twenty-two strains isolated 
from the placentas or fetuses of cows 
having abortion. 

Habitat: Causes abortion in cattle. 

22. Vibrio pierantonii (Zirpolo) Meiss- 
ner. {Bacillus pierantonii Zirpolo, Boll. 
Soc. nat. Napoli, 30, 1918, 206; Meissner, 
Cent. f. Bakt., II Abt., 67, 1926, 200.) 
Named for Pierantoni, an Italian bac- 
teriologist. 

Rods : 0.5 by 1.5 microns, with rounded 
ends. Motile with one to three polar 
flagella. Gram-negative. 

Gelatin colonies: Circular, and irregu- 
larly lobulate. 

Gelatin stab: No liquefaction. 

Agar colonies: Circular, light green, 
smooth, entire. 

Glycerin agar slant : Slightl}^ luminous 
streak. 

Broth : Turbid, with pellicle. 

Indole not formed. 

Acid from glucose and maltose. 
Some strains also attack lactose, sucrose 
and mannitol. 

Best growth in alkaline media. 

Aerobic, facultative. 

Optimum temperature 37 °C. 

Source : Isolated from the photogenic 
organ of the cephalopod Sepiola inter- 
media Naef . 

Appendix:* The following species have 
also been listed in the literature. Many 
are inadequately described. 



Microspira bonhojjli Migula. (Bonhoff , 
Arch. f. Hyg., 19, 1893, 252; Migula, 
Syst. d. Bakt., 2, 1900, 1008.) From 
water. 

Microspira canalis Migula. {Spiril- 
lum saprophiles y and Vibrio saprophiles 
7 Weibel, Cent. f. Bakt., 2, 1887, 469; 
Migula, Syst. d. Bakt., 2, 1900, 1004; 
Microspira cloaca Chester, Man. Determ. 
Bact., 1901, 341.) Possibly identical 
with Microspira saprophiles Migula, 
Microspira weibelii Migula, Vibrio siirati 
Ford, Vibrio smithii Ford. From sewage. 

Microspira coprophila Migula. (Group 
3, No. 6, Kutscher, Ztschr. f. Hyg., 19, 
1895, 475; Migula, Syst. der Bakt., 2, 
1900, 986.) From fecal matter. 

Microspira maasei (v. Hoff) Migula. 
{Spirillum maasei v. Hoff, Cent. f. Bakt., 
I Abt., 21, 1897, 797; Migula, Syst. d. 
Bakt., 2, 1900, 978.) Possibly a variety 
of Vibrio comma Winslow et al. From 
Rotterdam tap water. 

Microspira milleri Migula. (Miller, 
Deutsche med. Wchnschr., 11, 1885, 138; 
Migula, Syst. d. Bakt., 2, 1900, 981; 
Spirillum milleri Holland, Jour. Bact., 5, 
1920, 225; Vibrio milleri Holland, ibid.) 
Probably identical with Vibrio proteus 
according to Migula. From dental caries. 

Microspira rmirmanensis Issatchenko. 
(Recherches sur les microbes de I'ocean 
glacial arctique (in Russian), Petrograd, 
1914, 240.) From sea water. 

Microspira saprophiles Migula. (Heu- 
vibrio 13, Weibel, Cent. f. Bakt., 2, 1887, 
469 ; Vibrio saprophiles /3 Weibel, Cent, f . 
Bakt., 4, 1888, 225; Migula, Syst. d. 
Bakt., 2, 1900, 1006; Microspira weibell 
Chester, Man. Determ. Bact., 1901, 230.) 
Probably identical with Microspira cloaca 
Chester and Vibrio surati Ford. From 
sewage. 

Microspira tyrosinalica Beijerinck. 
(Kon. Akad. Wetenschappen, Amster- 
dam, 13, 1911, 1068.) From sewage. 

Microspira weibelii Migula. {Vibrio 



* Prepared by Mr. Wm. C. Ilaynes, New York State Experiment Station, Geneva, 
New York, Jan., 1939; Revised by Capt. Wm. C. Haynes, Sn. C, Fort Bliss, Texas, 
July, 1943. 



FAMILY PSEUDOMONADACEAE 



203 



saprophiles a Weibel, Cent. f. Bakt., 2, 
1887, 465; ibid., 4, 1888, 225; Migula, 
Syst. d. Bakt., 2, 1900, 1005; Microspira 
saprophilc Chester, Manual Determ. 
Bact., 1901,341; V ibrio saprophiles Ford, 
Textb. of Bakt., 1927, 356.) Possibly 
identical with Microspira cloaca Chester, 
Vibrio surati Ford. ]' ibrio smithii Ford. 
From sewage. 

Spirillum lipofcrutn Beijerinck. 
(Cent. f. Bakt., II Abt., 63, 1925, 353; 
Chromatium lipoferum Bergey et al., 
Manual, 3rd ed., 1930, 531.) From 
garden (>arth and sewage. Giesberger 
(Beitriige zur Kenntnis der Gattung 
Spirillum Ehbg., Inaug. Diss., Delft. 
1936, 64) regards this organism as a 
Vibrio. Has a single polar flagellum. 

SpiriUurn nasicola Trevisan. (Xasen- 
schleimvibrio, Weibel, Cent. f. Bakt., 
2, 1887, 465; Trevisan, I generi e le 
specie delle Batteriacee, 1889, 24; Vibrio 
nasalis Eisenberg, Bakt. Diag.. 3 Aufl., 
1891, 213; Spirillum nasale Sternberg, 
Man. of Bact., 1893, 697; Spirosoma 
nasale Migula, in Engler and Prantl, 
Die nattirl. Pflanzenfam., 1. I a, 1895. 
31.) From human nasal mucus. 

Spirillum parvum Esmarch. (Cent. f. 
Bakt., I Abt., Orig., 32, 1902, 565; also 
see Zettnow, ibid., 78. 1910, 1; Vibrio 
j'>arvus Lehmann and Neumann, Bakt. 
Diag., 4 Aufi., 2, 1907. 494.) From de- 
caying organic matter. 

Vibrio albensis Lehmann and Neu- 
mann. (Elbe vibrio, Dunbar, Deutsch. 
raed. Wochnschr., 19, 1893, 799; Leh- 
mann and Neumann, Bakt. Diag., 1 Aufl., 
2, 1896, 340; Microspira dunbari Migula, 
Syst. d. Bakt., 2, 1900, 1013; Photospiril- 
lum dunbari Miquel and Cambier, Traite 
de Bact., Paris, 1902, 881 ; Photobacterium 
dunbari Ford, Textb. of Bakt., 1927, 621 .) 
From water of the river Elbe. Phos- 
phorescent. 

Vibrio amylocella Gray. (Canad. Jour. 
Res., 17, 1939, 154.) Decomposes cel- 
lulose. Produces glucose from starch. 
From soil. 

Vibrio anguillarum Bergman. (Ber. a. 
d. k. Bayr. Biolog. Versuchstat., Mun- 



chen, 2, 1909.) From an infectious dis- 
ease of eels. 

Vibrio aureus Weibel. (Weibel, Cent, 
f. Bakt., 4, 18S8, 225, 257, 281; Spirillum 
aureum Trevisan, I generi e le specie 
delle Batteriacee, 1889, 24; Spirillum 
aureum Sternberg, Man. of Bact., 1893, 
700; Spirosoma aureum Migula, Syst. d. 
Bakt.. 2, 1900, 958.) Possibly identical 
with Vibrio flavns Weibel and Vibrio 
flavescens Weibel. From sewage. 

Vibrio beijerinckii Stanier. (Jour. 
Bact., 42, 1941, 527-554.) Marine agar- 
digesting vibrio. 

Vibrio buccalis Prevot. (Vibrion B, 
Repaci, Compt. rend. Soc. Biol., Paris, 
1909, 630; Prevot, :Man. de Classif. des 
Bact. Anacr., Paris, 1940. 82. i Anaerobe. 
From the buccal cavitJ^ 

Vibrio bulbosa Kalnins. (Latvijas 
Universitates Raksti, Serija I, No. 11, 
1930,237.) Decomposes cellulose. From 
soil. 

Vibrio cardii Klein. (Cent. f. Bakt., 
I Abt., Orig., 38, 1905, 173.) Possibly 
identical with Vibrio cuneatus Gray and 
Thornton and ]' ibrio marinus Ford. 
From the mussel (Cardium edule). 

Vibrio castra Kalnins. (Latvijas Uni- 
versitates Raksti, Serija I, No. 11, 1930, 
241.) Decomposes cellulose. From soil. 

Vibrio choleroides a and /3 Bujwid. 
(Cent. f. Bakt., 13, 1893, 120; Microspira 
choleroides ]\Iigula, Syst. d. Bakt., 2, 
1900, 992.) Probably a less vigorous 
strain of Vibrio comma Winslow et al. 
according to Chester, Man. Determ. 
Bact., 1901, 337. From water. 

Vibrio chrxjsanthemoides Lehmann and 
Neumann. (Spirillum-like organism, 
Jones. Cent. f. Bakt., II Abt.. 14. 1904, 
459; Lehmann and Neumann, Bakt. 
Diag., 4 Aufl., 2, 1907. 493.) From five 
samples of tap water and sewage. 

Vibrio crassus (Veillon and Repaci) 
Prevot. {Spirillum crassum Veillon and 
Repaci, Ann. Inst. Past., 26, 1912, 306; 
Prevot, Man. de Classif. des Bact. Anaer., 
Paris, 1940, 85.) Anaerobe. From the 
buccal cavity. 

Vibrio crassus var. D, Prevot. (Spirille 



204 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



D, Repaci, Ann. Inst. Past., 26, 1912, 
550; Prevot, Man. de Classif. des Bact. 
Anaer., Paris, 1940, 86.) Anaerobe. 
From the buccal cavity. 

Vibrio crasteri Hauduroy et al. 
(Craster, in Violle, Le Cholera, Masson 
edit., 1919; Hauduroy et al.. Diet. d. 
Bact. Path., 1937, 541.) Isolated from 
healthy persons. Resembles Vibrio 
comma. 

Vibrio cucumis Kalnins. (Latvijas 
tJniversi tales Raksti, Serija I, No. 11, 
1930,243.) Decomposes cellulose. From 
soil. 

Vibrio devorans Beijerinck. (Cent. f. 
Bakt., II Abt., 11, 1903, 598.) From 
water. 

Vibrio drennani Chalmers and Water- 
field. (Drennan, Jour. Inf. Dis., 14, 
1914, 251; Chalmers and Waterfield, 
Jour. Trop. Med., 19, 1916, 165.) Colon- 
ies white, turning dark brown. From 
feces. 

Vibrio flavescens Weibel. (Cent. f. 
Bakt., 4, 1888, 225, 257, 281; Spirillum 
flavescens Trevisan, I generi e le specie 
delle Batteriacee, 1889, 24; Spirillum, 
flavescens Sternberg, Man. of Bact., 1893, 
700; Spirosoma flavescens Migula, Syst. d. 
Bakt., 2, 1900, 959.) Possibly identical 
with Vibrio aureus Weibel and Vibrio 
flavus Weibel. From sewage. 

Vibrio flavus Weibel. (Cent. f. Bakt., 
4, 1888, 225, 257, 281; Spirillum flavum 
Trevisan, I generi e le specie delle Bat- 
teriacee, 1889, 24; Spirillum flavum 
Sternberg, Man. of Bact., 1893, 700; 
Spirosoma flavumM\g\i\&, Syst. d. Bakt., 
2, 1900, 959.) Possibly identical with 
Vibrio aureus Weibel and Vibrio flaves- 
cens Weibel. From sewage. 

Vibrio fuscus Stanier. (Jour. Bact., 
42, 1941, 540.) Marine agar-digesting 
vibrio. 

Vibrio gauducheau Hauduroy et al. 
(Gauducheau, in Violle, Le Cholera, 
Masson 6dit., 1919; Hauduroy et al., 
Diet. d. Bact., 1937, 543.) From the 
blood of a fever patient. Resembles 
Vibrio comma. 

Vibrio ghinda Pfeiffcr. (Pasquale, 



Gior. med. d. r. esercito, 1891 ; Pfeiffer, 
in Fliigge, Die Mikroorganismen, 2, 1896, 
590; Microspira ghinda Migula, Syst. d. 
Bakt., 2, 1900, 996.) From water. 

Vibrio grossus (Migula) Ford. (Vibrio 
No. 1, Kutscher, Ztschr. f. Hyg., 20, 
1895, 46 ; Microspira grossa Migula, Syst. 
d. Bakt., 2, 1900, 1012; Ford, Te.xtb. of 
Bact., 1927, 343.) From liquid manure. 

Vibrio halobicxis desulfuricans Horo- 
witz-Wlassowa and Sonntag. (Arb. a. 
d. Staatl. wissensch. Nahrungsmittel- 
Institut 1931 (Russian); see Ztschr. f. 
Unters. d. Lebensm., 62, 1931, 597.) A 
halophilic vibrio found in salted sar- 
dines, anchovies and other marine fish. 

Vibrio helcogenes Fischer. (Cent. f. 
Bakt., 14, 1894, 73; Microspira helcogenes 
Migula, Syst. d. Bakt., 2, 1900, 978.) 
From descriptions, indistinguishable 
from Vibrio proteus according to Chester, 
Man. Determ. Bact., 1901, 339. From 
feces. 

Vibrio hyos Ford. (Vibrio No. 3, 
Kutscher, Ztschr. f. Hyg., 20, 1895, 46; 
Spirillum mobile Migula, Syst. d. Bakt., 
2, 1900, 1020; Ford, Textb. of Bact., 1927, 
342.) Isolated from liquid manure. 

Vibrio iners Besson, Ranque and Senez. 
(Compt. rend. Soc. Biol. Paris, 79, 1918, 
1097.) From the feces of persons having 
dysentery. 

Vibrio intermedins (Migula) Ford. 
(Group V, No. 9 of cholera-like vibrios, 
Kutscher, Ztschr. f. Hyg., 20, 1895, 481; 
Microspira intermedia Migula, Syst. d. 
Bakt., 2, 1900, 967; Ford, Textb. of Bact., 
1927,342.) Possibly identical with Vib- 
rio berolinensis Neisser. From water. 

Vibrio ivanoff Pfeiffer. (Ivanoff, 
Ztschr. f. Hyg., 15, 1893, 134; Pfeiffer, 
in Fliigge, Die Mikroorganismen, 2, 1896, 
592.) Probably a variety of Vibrio 
comma Winslow et al. according to 
Chester, Man. Determ. Bact., 1901, 337. 
From feces of a cholera patient. 

Vibrio jejuni Jones, Orcutt and Little. 
(Jour. Exp. Med., 53, 1931, 853.) From 
small intestine of calves suffering from 
diarrhea. 

Vibrio l-egallcnsis Hauduroy et al. 



FAMILY PSEUDOMONADACEAE 



205 



(Diet. d. Bact. Path., 1937, 544.) From 
water. 

Vibrio klimenko Hauduroy et al. 
(Klimenko, in Violle, Le Cholera, Masson 
edit., 1919; Hauduroy et al., Diet. d. 
Bact. Path., 1937, 544.) Resembles 
Vibrio comma. From the intestine. 

Vibrio leidensis Horst. (Inaug. Diss., 
Leiden. 1921; abst. in Cent. f. Bakt., I 
Abt., Ref., 73, 1922, 282.) From a liver 
abscess. 

*Vibrio lingualis P^isenberg. (Zungen- 
belag-Vibrio, Weibel, Cent. f. Bakt., 4, 
1888, 227; ELsenberg. Bakt. Diag., 3 Aufl., 
1891, 212; Spirillum linguae Sternberg, 
Alan, of Bact., 1893, 697; Spirosoma lin- 
guale Aligula, in Engler and Prantl, Die 
naturl. Pflanzenfam.. /, 1 a, 1895, 31.) 
From deposit on the tongue. 

Vibrio lissabonensis Pestana-Betten- 
court. (Cent. f. Bakt., 16, 1894, 401.) 
According to Chantemesse identical, or 
nearly so, with Vibrio proleus. From 
descriptions, indistinguishable from Vib- 
rio proteus according to Chester, Man. 
Determ. Bact., 1901, 339. From feces 
of a cholera patient. 

Vibrio malamoria Kalnin.s. (Latvijas 
U^niversitates Raksti, Serija I, No. 11, 
'930,250.) Decomposes cellulose. From 
soil. 

Vibrio marinus (Russell) Ford. 
{Spirillum marinum Russell, Ztschr. f. 
Hyg., 11, 1891, 165; Microspira marina 
Migula, Syst. d. Bakt., 2, 1900, 1002; 
Ford, Textb. of Bact., 1927, 347.) From 
sea water. Closely resembles Vibrio 
cuneatus Gray and Thornton and Vibrio 
cardii Klein. 

Vibrio tuassauah Pfeiffer. (Pasquale, 
Gior. med. d. r. esercito, 1891; Pasquale, 
Baumgarten's Jahresberichte,7, 1891,336; 
Pfeiffer, in Flligge, Die Mikroorganismen, 
2, 1896, 589 ; Microspira massauah Migula, 
Syst. d. Bakt., 2, 1900, 963; Spirillum 
massauah Chester, Manual Determ. 
Bact., 1901, 343; Spirillum massowah 
Holland, Jour. Bact., 5, 1920, 225; Vibrio 
massowah Holland, ibid.) From feces 
of a cholera patient. 



Vibrio mulicris Prevot. (Man. de 
Classif. des Bact. Anaer., Paris, 1940, 84.) 
Anaerobe. From the female genital tract. 

Vibrio napi Kalninis. (Latvijas Uni- 
versitates Raksti, Serija I, No. 11, 1930, 
252.) Decomposes cellulose. From soil. 

Vibrio n'dianka Hauduroy et al. 
(Thiroux, in Violle, Le Cholera, Masson 
edit., 1919; Hauduroy et al.. Diet. d. 
Bact. Path., 1937, 546.) Isolated from a 
patient having a cholera-like disease. 

Vibrio pericoma Kalnins. (Latvijas 
tJniversitates Raksti, Serija I, No. 11, 
1930,256.) Decomposes cellulose. From 
soil. 

Vibrio polymorphus Prevot. (Spiro- 
chete B, Repaci, Ann. Inst. Past., 26, 
1912, 544; Vibrio pseudospirochaeta B, 
Weinberg, Nativelle and Prevot, Les 
Microbes Anaerobies, 1936, 852; Prevot, 
Man. de Classif. des Bact. Anaer., Paris, 
1940, 83.) Anaerobe. From the buccal 
cavity. 

Vibrio polymorphus var. peritriche 
Prevot. (Spirochete C, Repaci, Ann. 
Inst. Past., 26, 1912, 548; Vibrio pseudo- 
spirochaeta C, Weinberg, Nativelle and 
Prevot, Les Microbes Anaerobies, 1936, 
854; Prevot, Man. de Classif. des Bact. 
Anaer., Paris, 1940, 84.) Anaerobe. 
From the buccal cavity. 

Vibrio portuensis (Migula) Ford. (Der 
portuenser Vibrio, Jorge, Cent. f. Bakt., 
I Abt., 19, 1896, 277; Microspira portuen- 
sis Migula, Syst. d. Bakt., 2, 1900, 1007; 
Ford, Textb. of Bact., 1927, 353.) From 
water. 

Vibrio prima Kalnins. (Latvijas Uni- 
versitates Raksti, Serija I, No. 11, 1930, 
235.) Decomposes cellulose. From soil. 

Vibrio pseudospirochaeta Prevot. 
(Spirochete A, Repaci, Ann. Inst. Past., 
26, 1912, 539 ; Vibrio pseudospirochaeta A, 
Weinberg, Nativelle and Prevot, Les 
Microbes Anaerobies, 1936, 849; Prevot, 
Man. de Classif. des Bact. Anaer., Paris, 
1940, 83.) Anaerobe. From tlie buccal 
cavity. 

Vibrio putridus Prevot. (Vibrion C, 
Repaci, Compt. rend. Soc. Biol. Paris, 



See Nocardia lingualis Chalmers and Christopherson. 



206 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



1909, 630; Prevot, JMan. de Classif. des 
Bact. Anaer., Paris, 1940, 83.) Anaerobe. 
From the buccal cavity. 

Vibrio pyogenes (Doerr) Lehmann and 
Neumann. (Eiterspirillum, Mezinescu, 
Cent. f. Bakt., I Abt., Orig., 35, 1904, 
201; Spirillum pyogenes Doerr, Cent. f. 
Bakt., I Abt., Orig., S8, 1905, 15; Leh- 
mann and Neumann, Bakt. Diag., 4 
Aufl., 2, 1907, 493.) From pus in a case 
of pyelitis calculosa. Non-motile. 

Vibrio ranicula Kalnins. (Latvijas 
tjniversitates Raksti, Serija I, No. 11, 
1930,248.) Decomposes cellulose. From 
soil. 

Vibrio rigensis Kalnins. (Latvijas 
tjniversitates Raksti, Serija I, No. 11, 
1930,254.) Decomposes cellulose. From 
soil. 

Vibrio rubicundus Gottron et al. 
(Gottron, Weaver and Sherago, Jour. 
Bact., 4S, 1942, 61.) From a trickling 
filter. 

Vibrio septicus Kolle. (Kolle and 
Schumann in Kolle and Wassermann, 
Handb. d. path. Mikroorg., 2 Aufl., 4, 
1912, 101.) Identical with Vibrio comma 
culturally and morphologicalh*. From 
a cholera-like disease. 

Vibrio smithii (Migula) Ford. (Smith, 
Cent. f. Bakt., 10, 1891, 177; Microspira 
smithii Migula, Syst. d. Bakt., 2, 1900, 
1006; Ford, Textb. of Bact., 1927, 340.) 
Possibly identical v,-ith Microspira sapro- 
philes Migula, Microspira weibelii Mi- 
gula, Microspira cloaca Chester and 
Vibrio surati Ford. From abscesses of 
large intestine of swine. 

Vibrio spermatozoides Loffier. (Cent, 
f. Bakt., 7, 1890. 638.) From kohlrabi 
infusions. 

Vibrio sputigenus (Miller) Prevot. 
{Spirillum sputigenum Miller, Die Mikro- 
org. d. Mundhohle, 2nd ed., 1892; Prevot, 
Man. de Classif. des Bact. Anaer., Paris, 
1940, 85; not Vibrio sputigenus Bergey 
et al., Manual, 1st ed., 1923, 80.) An- 
aerobe. From the buccal cavity. 

Vibrio sputigenus var. minutissimus 
Prevot. (Muhlens, Cent. f. Bakt., I 



Abt., 48, 1909, 523; Pr6vot, Man. de 
Classif. des Bact. Anaer., Paris, 1940, 85.) 
Anaerobe. From the buccal cavity. 

Vibrio sputorum Prevot. (Man. de 
Classif. des Bact. Ana6r., Paris, 1940, 85.) 
Anaerobe. Isolated from a case of 
bronchitis. 

Vibrio stationis Kalnins. (Latvijas 
tjniversitates Raksti, Serija I, No. 11, 
1930,239.) Decomposes cellulose. From 
soil. 

Vibrio stomatitis Prevot. (Vibrion A, 
Repaci, Compt. rend. Soc. Biol. Paris, 
1909, 630; Prevot, Man. de Classif. des 
Bact. Anaer., Paris, 1940, 82.) Anaerobe. 
From the buccal cavity. 

Vibrio subtilissimus (Migula) Ford. 
(Spirillum No. 1, Kutscher, Ztschr. f. 
Hyg., 20, 1895, 46; Spirillum tenerrimum 
Lehmann and Neuman, Bakt. Diag., 2, 
1896, ZA^; Spirillum subtilissimumMigula, 
Syst. d. Bakt., 2, 1900, 1020; Ford, Textb. 
of Bact., 1927, 341.) Regarded by 
Kutscher as being probably identical 
with the organism found by Smith (Cent, 
f. Bakt., 16, 1894, 324) in swine dung. 
Resembles Vibrio strictus. 

Vibrio suis Ford. (Vibrio No. 2, 
Kutscher, Ztschr. f. Hyg., 20, 1895, 46; 
Spirillum coprophilum Migula, Syst. d. 
Bakt., 2, 1900, 1019; not Microspira 
coprophila Migula, loc. cit., 986; Ford, 
Textb. of Bact., 1927, 341.) From liquid 
manure. 

Vibrio surati (Lamb and Baton) Ford. 
{Spirillum surati Lamb and Paton, Arch. 
Int. Med., 12, 1913, 259; Treponema 
surati Brumpt, Nouveau Traite de 
Medecine, Paris, 4, 1922, 514; Ford, 
Textb. of Bact., 1927, 337.) Isolated 
from a case of vegetative endocarditis. 
Closely resembles Vibrio smithii Ford, 
Microspira weibelii Migula, Microspira 
saprophiles Migula and Microspira cloaca 
Chester. 

Vibrio synthetica Kalnins. (Latvijas 
tjniversitates Raksti, Serija I, No. 11, 
1930,245.) Decomposes cellulose. From 
soil. 

Vibrio tenuis Veillon and Repaci. 



FAMILY PSEUDOMONADACEAE 



207 



(Ann. Inst. Past., 26, 1912, 300.) An- 
aerobe. From the buccal cavity. 

Vibrio terrigenus Giinther. (Cent. f. 
Bakt., 16, 1894, 746; Spirillum terri- 
genum Migula, Syst. d. Bakt., 2, 1900, 
1017; Microspira terrigena Chester, Man. 
Determ. Bact., 1901, 341.) Closely re- 
lated to Vibrio tonsillaris Stephens and 
Smith. From soil. 

Vibrio tonsillaris Stephens and Smith. 
(Cent. f. Bakt., 19, 1896, 929; Microspira 
tonsillaris Migula, Syst. d. Bakt., 2, 



1900, 1009.) Closely related to Vibrio 
terrigenus Giinther. From buccal cavity. 

Vibrio toulonensis Hauduroy et al. 
(Vibrion, Defressine and Cazeneuve, in 
Violle, Le Cholera, Masson edit., 1919; 
Hauduroy et al.. Diet. d. Bact. Path., 
1937, 547.) From mussel beds in the 
bay of Toulon. 

Vibrio xylitica Kalnins. (Latvijas 
tJniversitates Raksti, Serija I, No. 11, 
1930,232.) Decomposes cellulose. From 
soil. 



Genus II. Desulfovibrio Kluyver and van Niel.* 

(Cent. f. Bakt., II Abt., 04, 1936, 369; Sporovibrio Starkey, Arch. f. Mikrobiol., 9, 

1938, 300.) From M. L. desulfo, an abbreviation of the poorly constructed word 

desulfofication, used to indicate reduction of sulfur compounds by bacteria; 

vibrio, vibrio. 

Slightly curved rods of variable length, usually occurring singly but sometimes in 
short chains which have the appearance of spirilla. Swollen pleomorphic forms are 
common. Actively motile by means of a single polar flagellum. Strict anaerobes 
which reduce sulfates to hydrogen sulfide. Found in sea water, marine mud, fresh 
water, and soil. 

The type species is Desulfovibrio desulfuricans (Beijerinck) Kluyver and van Niel. 



1. Desulfovibrio desulfuricans 
(Beijerinck) Kluyver and van Niel. 
{Bacterium hydrosuljureum ponticum 
Zelinsky, Proc. Russ. Phys. and Chem. 
Soc, 25, 1893, 298; Spirillum desul- 
furicans Beijerinck, Cent. f. Bakt., II 
Abt., 1, 1895, 1: Bacillus desidfuricans 
Saltet, Cent. f. Bakt., II Abt., 6, 1900, 
648; Microspira desulfuricans Migula, 
Syst. d. Bakt., 2, 1900, 1016; Kluyver 
and van Niel, Cent. f. Bakt., II Abt., 
94, 1936, 369; Vibrio desulfuricans 
Holland, Jour. Bact., 5, 1920, 225; Sporo- 
vibrio desulfuricans Starkey, Koninkl. 
Nederland. Akad. v. Wetenschappen, 
Proc, 41, 1938, 425; also in Arch. f. 
Microbiol., 9, 19.38, 268.) From U. L. 
present part, desidfurico, sulfur re- 
ducing. 

Slightly curved rods, 0.5 to 1.0 by 1 to 5 
microns, usually occurring singly but 
sometimes in pairs and short chains 



which cause them to look like spirilla. 
Swollen pleomorphic forms are common. 
Older cells appear black due to precipi- 
tated ferric sulfide. Actively motile, 
possessing a polar flagellum. Gram- 
negative. Stains readilj' with carbol 
fuchsin. 

Grows best in freshwater media. 
Fails to develop in sea water upon initial 
isolation. 

Produces opalescent turbiditj' in ab- 
sence of oxygen in mineral media enriched 
with sulfate and peptone. 

Media containing iron salts blackened. 
Bacteria found associated with pre- 
cipitated ferrous sulfide. 

Peptone-glucose agar colonies (in 
absence of air) : Small, circular, slightly 
raised, dull, entire, soft in consistency. 

Gelatin not liquefied. 

Peptone, asparagine, glycine, alanine, 
aspartic acid, ethanol, propanol, butanol, 



* Prepared by Dr. Claude E. 
.Jolla, California, Jan., 1943. 



ZoBell, Scripps Institution of Oceanography, La 



208 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



glycerol, glucose, lactate, succinate and 
malate known to be utilized as hydrogen 
donors. 

Produces up to 500 ml. H2S per liter. 

Nitrites not produced from nitrates. 

Reduces sulfate to hydrogen sulfide. 
Also reduces sulfites, sulfur, thiosulfates 
and hyposulfites. 

Optimum pll 6 to 7.5, limits pH 5 to 9. 

Optimum temperature 25 to 30° C. 
Maximum 35 to 40° C. 

Anaerobic. 

Habitat: Soil, sewage, water. 



aestuarii (van 
nov. {Microspira 



2. Desulfovibrio 
Delden) comb, 
aestuarii van Delden, Cent. f. Bakt., II 
Abt., 11, 1904, 81; Vibrio dcsulfur leans 
(halophilic strain) Baars, Over Sulfaat- 
reductie door Bakterien, Diss. Delft, 
1930, 164 pp.) From Latin, aestuarium, 
estuary. 

Morphologically indistinguishable from 
Desulfovibrio desulfur leans described 
above, although it has a greater tendency 
to pleomorphism, and is slightly larger. 
Motile, possessing a polar flagellum. 
Gram-negative. 

Grows preferentially in media prepared 
with sea water or 3 per cent salt mineral 
solution enriched with sulfate and 
peptone. According to Baars {loc. clt.) 
the marine species can be acclimatized to 
tolerate hypotonic salt solutions but 
Rittenberg (Studies on Marine Sulphate- 
Reducing Bacteria, Thesis, Univ. of 
Calif., 1941, 115 pp.) was unable to con- 
firm this observation. Likewise Ritten- 
berg was unable to acclimatize D. aes- 
tuarii to tolerate temperatures exceeding 
45° C or to produce endospores. 

Produces faint turbidity in absence of 
oxygen in sea water enriched with sulfate 
and peptone. Organisms most abundant 
in sediment. 

Agar colonies: Small, circular, slightly 
raised, darker centers, entire, soft 
consistency. 

Gelatin not liquefied. 

Peptone, asparagine, glycine, alanine, 
glucose, fructose, ethanol, butanol. 



glycerol, acetate, lactate and malate 
known to be utilized in presence of 
sulfate. 

Reduces sulfate to hydrogen sulfide. 
Also reduces sulfites, sulfur, thiosulfates 
and hyposulfites. 

Produces up to 950 ml. H2S per liter. 

Nitrites not produced from nitrates. 

Optimum temperature 25° to 30° C. 
Maximum 35° to 40° C. 

Optimum pH 6 to 8, limits pH 5.5 to 
8.5. 

Anaerobic. 

Habitat : Sea water, marine mud, brine 
and oil wells. 

3. Desulfovibrio rubentschickii 

(Baars) eomb. nov. {Vibrio rubent- 
schickii Baars, Over Sulfaatreductie 
door Bakterien, Diss. Delft, 1930, 164 
pp.) Named for L. Rubentschick. 

Slightly curved rods, 0.5 to 1.0 by 
1 to 5 microns, usually occurring singly, 
sometimes in pairs and short chains. 
Actively motile, possessing a polar 
flagellum. Gram-negative. Morpho- 

logically indistinguishable from Desul- 
fovibrio desulfuricans. 

Reduces sulfate to hydrogen sulfide. 
Also reduces sulfites, sulfur, thiosulfates 
and hyposulfites. 

Culturally and physiologically like 
D. desulfuricans except that D. rubent- 
schickii utilizes propionic acid, butyric 
acid, valeric acid, palmitic acid, stearic 
acid, galactose, sucrose, lactose and 
maltose. 

Anaerobic. 

Habitat : Soil and ditch water. 

Appendix : The following species has 
also been regarded as belonging in this 
genus. 

Vibrio thermodesulfurlcans Elion. 
(Cent. f. Bakt., II Abt., 63, 1924, 58); 
Vibrio desulfuricans (thermophilic 
strain) Baars, Over Sulfaatreductie door 
Bakterien, Diss. Delft, 1930, 164 pp.; 
Sporovibrio desulfuricans Starkey (Ko- 
ninkl. Nederland. Akad. u. Wetenschap- 
pen, Proc, 41, 1938, 425, also see Arch. f. 



FAMILY PSEUDOMONADACEAE 209 

Microbiol., 5, 1938, 268.) A thermophilic produces endospores. However, spore- 

sulfate-reducing anaerobe which grows at formation appears to be the exception 

30 to 65°C. and which, according to rather than the rule. The pleomorphic, 

Starkey, produces endospores. Elion peritrichous, sporogenous, sulfate- 

described Vibrio thcrmodesulfuricans reducer is more rod-like than the asporo- 

(Cent. f. Bakt., II Abt., 63, 1924, 58) genous cultures and many cells of the 

which grows at temperatures no lower sporogenous cultures are Gram-positive 

than 30 to 40°C. and has an optimum of ^^.^^^^^^^ asporogenous cultures of Desnl- 

55°C. Morphologically it is much like r -i ■ i u- ■ r^ j.- 

^ ,^ , , ,,. • , rx f ovibr 10 desulfuncans are {jvam-negative, 

Desulfoinbrio desulfuricans and D. aes- " n ,• , ■ > , ,. , ,, ^i 

.. , ,, , ., ,, , ■,- r • all oi which leaves a question whether the 
iuarii although the thermophilic form is ,r , • r. -77 

, , J 1-1 1 i-i I sporogenous suliate-reducer is a Bacillus 

shorter, more rod-like, less motile and 

more pleomorphic. According to Baars «^' ^^ Desulfoinbrto. Rittenberg (htudies 
(loc. cit.), Vibrio thermodesulfuricam »" ^^^^"^e Sulfate-reducing Bacteria, 
Elion can be acclimatized to grow at Thesis, Univ. Calif., 1941, 115 pp.) was 
lower temperatures and it is found unable to adapt the marine sulfate re- 
abundantly in environments where the ducer to grow at low salinities or at high 
temperature has never been as high as temperatures, nor could it be induced 
30 °C. This observation is confirmed by to form spores. 

Starkey (Arch. f. Microbiol., 9, 1938, Desulforibrio halohydrocarbonoclasti- 

268) who found further that the thermo- cus Zobell (U. S. Patent Xo. 2.413,278; 

philic form found in nature or developed Science News Letter, Jan. 1 1, 1947.) 

b}' acclimatization to higher temperatures From oil bearing rocks. 

Genus III. Cellvibrio Winogradshy* 
(Ann. Inst. Pasteur, 4^, 1929. 577.) From M. L. cell, an abbreviation for cellulose; 

vibrio, vibrio. 
Long slender rods, slightly curved, with rounded ends, show deeply staining gran- 
ules which appear to be concerned in reproduction. Monotrichous. Most species 
produce a yellow or brown pigment with cellulose. Oxidize cellulose, forming oxy- 
cellulose. Growth on ordinary culture media is feeble. Found in soil. 
The type species is Cellvibrio ochraceus Winogradsky. 

Key to the species of genus Cellvibrio. 
I. Xo growth on glucose or starch agar. 

A. Ochre-yellow pigment produced on filter paper. 

1. Cellvibrio ochraceus. 
II. Growth on glucose and starch agar. 

A. Poor growth on starch agar. 

1. Cream-colored pigment which becomes brown with age is produced on 
filter paper. 

2. Cellvibrio flavescens. 

B. Abundant growth on starch agar. 

1. Scanty growth on glucose agar. 

a. Intense yellow pigment produced on filter paper. 

3. Cellvibrio fulvus. 

2. Abundant growth on glucose agar. 

a. X"o pigment produced on filter paper. 

4. Cellvibrio vulgaris. 

* Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva, 
New York, Sept., 1937; no change. July, 1943. 



210 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



1. Cellvibrio ochraceus Winogradsky. 
(Ann. Inst. Pasteur, J!^3, 1929, 549, 601.) 
From Greek, ochra, yellow ochre; M. L. 
like ochre, yellow. 

Plump, curved rods with rounded 
ends, 2.0 to 4.0 microns long, rarely 
occurring as spirals. Chromatic granule 
frequently found in center. Motile 
with a single flagellum. Gram-negative. 

Produces diffuse, light ochre-colored, 
mucilaginous colonies on cellulose silica 
gel medium. 

No action or growth on plain agar. 
No growth on peptone, glucose, starch 
or tragacanth gum agar. 

Filter paper streaks : Entire paper 
colored ochre-yellow in 48 hrs. 

Aerobic, facultative. 

Optimum temperature 20 °C. 

Distinctive character : Rapid ochre- 
colored growth. 

Habitat : Soil. Disintegrates vegetable 
fibers. 

2. Cellvibrio flavescens Winogradsky. 
(Ann. Inst. Pasteur, 43, 1929, 608.) From 
Latin, part. adj. oi flavesco, to turn yel- 
low or golden. 

Plump, curved rods, flexuous, with 
rounded ends, 0.5 by 2.5 to 5.0 microns. 
Shows metachromatic granules. Motile 
with a single flagellum. Gram-negative. 

Produces diffuse, cream-colored growth 
becoming brownish; mucilaginous colo- 
nies on cellulose silica gel medium. 

Good growth on peptone agar. Colo- 
nies 1 mm in 4 days. Grows poorly on 
glucose, starch and gum agars. 

Filter paper streaks : Almost as rapid 
in growth as Cellvibrio ochraceus and 
colors entire paper in 2 to 3 days. 

Aerobic, facultative. 

Optimum temperature 20 °C. 

Distinctive characters : Smaller, less 
curved rods that grow on a greater 
variety of media than Cellvibrio ochra- 
ceus, but do not attack cellulose as 
readily. 

Source : Isolated from a pile of old 
damp sawdust. 



Habitat : Soil. Disintegrates vegetable 
fibers. 

3. Cellvibrio fulvus Stapp and Bortels. 
(Culture Y, Dubos, Jour. Bact., 15, 
1928, 230; Stapp and Bortels, Cent. f. 
Bakt., II Abt., 90, 1934, 42.) From 
Latin, fulvus, reddish yellow. 

Slightly curved rods: 0.3 to 0.4 by 
1.5 to 3.0 microns. Show involution 
forms. Motile by means of a single polar 
flagellum. Gram-negative. 

Cellulose is decomposed. Grows on 
filter paper with an intense egg-yellow 
color which in older cultures may deepen 
to rust brown. 

Glucose agar: Very scanty growth. 

Sucrose agar: Very slight growth. 

Maltose agar : Abundant yellow growth. 

Lactose agar: Fairly abundant yellow 
growth. 

Starch agar: Very abundant, bright 
yellow growth which later turns brown. 

Nutrient broth : No growth. 

Temperature relations: Optimum 25° 
to 30°C. Minimum 5°C. Maximum 32° 
to 35 °C. No growth at 37 °C. Thermal 
death point 39° to 40 °C. 

Aerobic. 

Source : Isolated from forest soil in 
Germany and from soil in the United 
States. 

Habitat : Widely distributed in soils. 

4. Cellvibrio vulgaris Stapp and 
Bortels. (Culture Co, Dubos, Jour. 
Bact., 15, 1928, 230; Stapp and Bortels, 
Cent. f. Bakt., II Abt., 90, 1934, 44.) 
From Latin, vulgaris, common. 

Curved rods : 0.3 by 2.9 to 4.0 microns. 
Shows involution forms. Motile by 
means of a single polar flagellum. Gram- 
negative. 

Cellulose is decomposed. Grows on 
filter paper without the formation of 
pigment. 

Glucose agar: Abundant growth. No 
pigment. 

Sucrose agar : Abundant slightly yellow 
growth. 



FAMILY PSEUDOMONADACEAE 



211 



Maltose agar: Abunilant yellowish to 35°C. No growth at 37 °C. Thermal 

growth. death point 44° to 45°C. 

Lactose agar: Very heavy growth. Aerobic 

Starch agar : Very abundant yellowish „ t , , , 

, , bource : Isolated from forest soil in 
growth . 

Nutrient broth : No growth. Germany and from soils in the United 

Temperature relations: Optimum 25° states. 

to30°C. Minimum 5°C. Maximum 32° Habitat: Widely distributed in soils. 



Genus IV. Cellfalcicula W inogradshy .* 

(Ann. Inst. Pasteur, 43, 1929, 616.) From M. L. cell, an abbreviation for cellulose; 
Latin dim.., Jalcicula, a small sickle. 

Short rods or spindles, not exceeding 2.0 microns in length, with pointed ends, 
containing metachromatic granules. Old cultures show coccoid forms. Monotri- 
chous. Oxidize cellulose, forming o.xycellulose. Growth on ordinary culture media 
is feeble. Soil bacteria. 

The type species is Cellfalcicula viridis Winogradsky. 



1. Cellfalcicula viridis Winogradsky. 
(Ann. Inst. Pasteur, iS. 1929. 616 .) From 
Latin, viridis, green. 

Plump, small spindles, 0.7 by 2.0 
microns, with rounded ends. Motile 
with a single flagellum. Gram-negative. 

Produces diffuse green, mucilaginous 
colonies on cellulose silica gel medium. 

Filter paper streaks : Rapid spreading 
growth colored green in 3 days at 30 °C. 

Hydrocellulose agar: Growth rapid, 
green; minute yellowish-green, mucous 
colonies on streaking. 

No growth on peptone, glucose, starch 
or gum agar. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Habitat : Soil. 

2. Cellfalcicula mucosa Winogradsky. 
(Ann. Inst. Pasteur, 43, 1929, 621.) From 
Latin, viucosus, mucous. 

Plump, curved spindles, with slightly 
pointed ends. Motile with a single polar 
flagellum. Contain a single chromatic 
granule. Gram-negative. 

Produces diffuse, cream-colored, muci- 



laginous colonies on cellulose silica gel 
medium. 

Hydrocellulose agar : Abundant grayish 
growth. 

No growth on peptone, glucose, 
starch or gum agar. 

Aerobic, facultative. 

Optimum temperature 20°C. 

Habitat: Soil. 

3. Cellfalcicula fusca Winogradsky. 
(Ann. Inst. Pasteur, 43, 1929, 622.) From 
Latin, fuscus, dark, tawny. 

Plump, curved spindles, 0.5 by 1.2 to 
2.5 microns, with slightly pointed ends 
and a central chromatic granule. Motile 
with a single polar flagellum. Gram- 
negative. 

Produces diffuse, brownish, slightly 
marbled or veined colonies on cellulose 
silica gel medium. 

Filter paper streak : Paper becomes a 
partially transparent, dry, non-muci- 
laginous pellicle adhei-ent to gel. 

Aerobic, facultative. 

Optimum temperature 20 °C. 

Source : Isolated from a pile of old damp 
sawdust. 

Habitat: Probably rotting wood. 



* Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva 
i^ew \ork, Sept., 1937; no change, July, 1943. 



212 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Genus V. Thiospira Vislouch-* 

(Jour, de Microbiologie, 1, 1914, 50; Sidjospir ilium Kluyver and van Niel, Cent. f. 
Bakt., II Abt., 9J^, 1936, 396.) From Greek, theion, sulfur; speira, coil. 

Colorless, motile, slightly bent rods, somewhat pointed at the ends, with granules 
of sulfur within the cells and a small number of flagella at the ends. 

The type species is Thinspira winograchkyi (Omelianski) Vislouch. 

1. Thiospira winogradskyi (Ome- 2. Thiospira bipunctata (Molisch) 

lianski) Vislouch. {ThiospiriUum wi7io- Vislouch. (Spirillum bipunctatum Mo- 

gradskyi Omelianski, Cent. f. Bakt., II Hsch, Cent. f. Bakt., II Abt., 33, 1912, 

Abt., 14, 1905, 7m ; ThiospiriUum. granu- 55; Vislouch, Jour, de Microbiologie 

latum Molisch, Cent. f. Bakt., II Abt., (Russian), 1, 1914, 50.) From Latin, bi, 

33, 1912, 55; Vislouch, Jour, de Micro- two; punctum, points, 

biologic '(Russian), 1, 1914, 50; Sulfo- Small, slightly bent sulfur spirilla, 

spirillum winogradskyi Kluyver and van markedly pointed at the ends, 6.6 by 14 

Niel, Cent. f. Bakt., II Abt., 94, microns long 1.7 to 2 4 microns wide (in 

,„„„ „„_ , T-T J i- ^^T■ J 1 the center of the cell). Both ends are 

1936, 397.) Named for Wmogradsky, -xi i i i- 

^ . , . , . , filled more or less with large volutin 
the Russian bacteriologist. (metachromatic) granules. Several mi- 
Large, sulfur spirilla, somewhat ^^^^^ granules of sulfur are present in the 
pointed at the ends, 2 to 2.5 microns ^j^^^. ^^^^^^ ^^^ sometimes at the ends, 
thick, to 50 microns long. Numerous qjj ^gjjg possess one flagellum at each 
granules of sulfur. Very motile, with end; young cells have a flagellum at 
one to two polar flagella. one end. 

Habitat: Curative mud. Habitat: Sea and salt waters. 

Gcnvs VJ. Spirillum Ehrenberg .^ 

(Ehrenberg, Abhaudlungon d. Berl. Akad., 1830, 38; Spirosoma Migula, Arb. bakt. 

Inst. Karlsruhe, 1, 1894, 237; Dicrospirillum Enderlein, Sitzber. Gesell. naturf. 

Freunde, Berlin, 1917, 313.) From Greek, speira, a spire or coil. 

Cells form either long screws or portions of a turn. Volutin granules are usually 
present. Usually motile by means of a tuft of polar flagella (5-20) which may occur 
at one or both ends of the cells. Aerobic, growing well on ordinary culturemedia, 
except for one saprophyte and the pathogenic species. These have not yet been 
cultivated. Usually found in fresh and salt water containing organic matter. 

The type species is Spirillum xindula (Miiller) Ehrenberg. 

Key to the species of genus Spirillum. 

I. One micron or less in diameter. 
1. Volutin granules present. 

a. Slow to rapid liquefaction of gelatin, 
b. Grayish to brown growth on potato. 

1. Spirillum vndula. 

* Prepared by Prof. D. H. Bergey, Philadelphia, Penn., October, 1922. 

t Revised by Prof. D. H. Bergey, Philadelphia, Pennsylvania, April, 1937; further 
revision by Prof. Robert S. Breed, New York State E.xperiment Station, Geneva, 
New York, June, 1943, based on Monograph by Giesberger, Inaug. Diss., Utrecht, 
Nov. 30, 1936. 



FAMILY PSEUDOMOiVADACEAE 



213 



bb. Light yellowish-orange growth on potato. 

2. Spirillum serpens. 

aa. No liquefaction of gelatin. Of small size (0.5 micron in diameter). 
b. Colonies on agar white becoming brownish black and slightly 
wrinkled. 

3. Spirillum itersonii. 
bb. Colonies on agar white and smooth. 

4. Spirillum tenue. 
2. No volutin granules observed. 

b. Single flagellum. 

5. Spirillum virginianum. 
bb. Tuft of flagella. 

6. Spirillum minus. 
II. Over one micron in diameter. 

1. Grows poorly on peptone agar and potato. 

7. Spirillum kutscheri. 

2. Xot positively known to have been cultivated on artificial media. Very 
evident volutin granules. 

S. Spirillum volulans. 

3. Cells more or less deformed by fat drops. 

9. Spirillum lipoferum. 



1. Spirillum undula rMiiller) Ehren- 
berg. {Vibrio undula MtiUer, Animal- 
cula infusoria et marina, 1786 ; Ehrenberg, 
Infusionstierchen, 1838; Spirillum 
undula minor Kutscher, Cent. f. Bakt., 
I Abt., 18, 1895, 614.) From Latin, 
undulatus, wave-like. 

Stout threads, 0.9 micron in diameter, 
with one-half to three turns. The wave 
lengths are 6 microns. Width of spiral, 
3.0 microns. Tufts of three to nine 
flagella at each pole. Volutin granules 
present. Gram-negative. 

Gelatin colonies : The surface colonies 
are circular, granular, greenish-yellow, 
entire. 

Gelatin stab: Thick, white, rugose 
surface growth. Very slow liquefaction. 

Agar colonies: Grayish-white, smooth. 

Broth : Turbid. 

Potato: Grayish-brown growth. 

Indole not formed. 

Catalase positive. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 25 °C. 

Cohn (Beitrage z. Biol. d. Pflanzen, /, 
Heft 2. 1875, 132) reports that he could 



not distinguish this organism from Vibrio 
prolifer Ehrenberg. 
Habitat: Putrid and stagnant water. 

2. Spirillum serpens (MuUer) Winter. 
{Vibrio serpens Miiller, Animalcula in- 
fusoria et marina, 1786, 43; Winter, in 
Rabenhorst's Kryptogamen-Flora, 1, Die 
Pilze, 1884, 63.) From Latin, serpens, 
serpent. 

Long, curved rods with two to three 
wave-like undulations, 0.8 to 1.0 micron 
in diameter; wave length, 8 to 9 microns. 
Width of spiral 1.5 to 1.8 microns. Vo- 
lutin granules in cytoplasm. Motile, 
possessing tufts of flagella at both poles. 
Gram-negative. 

Gelatin colonies : Yellowish to brown- 
ish, granular, entire. 

Gelatin stab : Yellowish surface growth. 
Slow liquefaction. 

Agar colonies : Heavj- cream-colored 
growth. 

Agar slant: Grayish, with yellowish 
center, granular, entire. 

Broth: Turbid. 

Litmus milk : Unchanged. 

Potato: Clear orange-yellow growth 

Indole not formed. 



214 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Catalase positive. 

Nitrites not produced from nitrates. 

Aerobic, facultative. 

Optimum temperature 35 °C. 

Habitat : Stagnant water. 

3. Spirillum itersonii Giesberger. 
(Inaug. Diss., Utrecht, 1936, 46 and 57.) 
Named for van Iterson, the Dutch bac- 
teriologist. 

The smallest of the spirilla isolated 
from water. First observed by van 
Iterson (Proc. Kon. Akad. v. Wetensch. 
Amsterdam, 5, 1902, 685). 

Small spirals, 0.5 micron in diameter. 
Wave length, 3 to 3.5 microns. Spiral 
width, 1 to 1.5 microns. Motile with 
bipolar tufts of flagella. Gram-negative. 

Grows readily on peptone agar. White 
colonies becoming brownish black, and 
slightly wrinkled. 

Gelatin stab : No liquefaction. 

Brownish-orange growth on potato. 

Volutin granules may be present. 

Catalase is i)roduced. 

Acid from glucose, fructose, ethyl 
alcohol, n-propyl alcohol, n-butyl alcohol, 
and glycerol. Utilizes acetic, propionic, 
n-butyric, tartaric, fumaric, lactic, citric, 
and succinic acids. 

Grows well in peptone broth. Also 
utilizes ammonia compounds. 

Anaerobic growth in the presence of 
nitrates when organic or ammonia nitro- 
gen is also available. 

Optimum temperature : 30°C. 

Source : Isolated from water. 

Habitat: Water. 

4. Spirillxun tenue Ehrenberg. (In- 
fusionstierchen, 1838; see Bonhoff, Arch, 
f. Hyg., ^6, 1896, 162.) From Latin, 
tenuis, thin. 

Slender spirals. Diameter 0.7 micron. 
Wave lengths 4.5 to 5.0 microns. Width 
of spiral 1.5 to 1.8 microns. 

Actively motile in peptone water with 
tufts of flagella at each pole. Volutin 
granules present. Gram-negative. 

Agar colonies: White, smooth. 



Peptone agar slant : Heavy growth. 

Gelatin stab : No liquefaction. 

Catalase positive. 

Potato : Light brown growth. 

Acid from glucose and fructose. 
Slight acid from several other sugars and 
glycerols. Utilizes salts of acetic, pro- 
pionic, n-butyric, tartaric, lactic, citric, 
malic, and succinic acids. 

Ammonia compounds are used as a 
source of nitrogen. 

Optimum temperature, 30 °C. 

Source : Found in putrefying vegetable 
matter. 

Habitat: Putrefying materials. 

5. Spirillum virginianum Dimitroff. 
(Jour, of Baet., 1:2, 1926, 19.) From M. 
L. genitive of Virginia. 

Spirals consisting of J to 3 complete 
turns in young cultures, older cultures 
showing 7 turns. 0.6 to 0.9 by 3 to 11 
microns. Motile with a single polar 
flagellum on one or both ends. Gram- 
negative. 

Gelatin colonies: Entire, convex, cir- 
cular, moist, colorless. 

Gelatin stab : Growth along entire stab. 
No liquefaction. (Dimitroft", loc. cit.) 
Active liquefaction. (Giesberger, Inaug. 
Diss., Utrecht, 1936, 65.) 

Agar colonies: Dew drop, convex, 
entire, moist, colorless. 

Agar slant : Dew drop, isolated colonies. 

Broth: Cloudy, no flocculation. 

Uschinsky's protein-free medium : 
Abundant growth. 

Litmus milk: No growth. 

Loeffler's blood serum : Convex, 
isolated dew drop colonies. No lique- 
faction. 

Lead acetate agar: No H2S. 

Voges-Proskauer and methyl red nega- 
tive. 

No volutin granules observed (Gies- 
berger, loc. cit., p. 60). 

Potato: No growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

No acid or gas from carbohydrates.- 



FAMILY PSEUDOMONADACEAE 



215 



(Dimitroff, loc. cit.). Utilizes lactates 
and citrates (Giesberger, loc. cit.). 

Aerobic, facultative. 

Optimum temperature 35 °C. 

Source : Isolated from mud on an oyster 
shell. 

Habitat : Probably muddy bottom of 
brackish water. 

6. Spirillum minus Carter. (Carter, 
Sci. Mem. Med. Officers Army India, 3, 
1887, 45; Spirillum minor Carter, ibid.; 
Spirochaeta laverani Breinl and King- 
horn, Mem. Liverpool Sch. Trop. Med., 
21, 1906, 55; Spirochaeta miiri.'i Wenyon, 
Jour. Hyg., 6, 1906, 580; Spirochaeta 
muris var. virginiana MacNeal, Proc. 
Soc. Exper. Biol, and Med., 4, 1907, 125; 
Spirochaeta muris var. galatziana Mezin- 
cescu, Compt. rend. Soc. Biol. Paris, 66, 
1909, 58; Treponema muris Moore, 
Principles of Microbiology, 1912, 414; 
Spirochaeta morsus mnris Futaki, Takaki, 
Taniguchi and Osumi, Jour. Exp. Med., 
25, 1917, 33; Spirochaeta petit Row, Ind. 
Jour. Med. Res., 5, 1917, 386; Spironema 
muris Noguchi, Jour. Exp. Med., 27, 

1918, 584; Spirochaeta japonica, Dujarric 
de la Riviere, Ann. de Med., 5, 1918, 184; 
Spirochaeta morsusmuris Castellani and 
Chalmers, Man. Trop. Med., 3rd ed., 

1919, 447; Spiroschaudinnia morsusmuris 
Castellani and Chalmers, ibid.; Spiro- 
chaeta sodoku Troisier, 1920, according to 
Pettit, Contribution a I'Etude des 
Spirochetides, Vanves, II, 1928, 231; 
Treponema japonicum Brumpt, Nouveau 
Traitc de Medecine, Paris, 4, 1922, 505; 
Treponema morsus muris Brumpt, ibid., 
506; Treponema minor Brumpt, ibid., 
507; Treponema laverani Brumpt, ibid., 
507; Treponema sodoku Brumpt, ibid., 
514; Spirochaeta pettiti Row, Jour. 
Trop. Med. and Hyg., 25, 1922, 364; 
Treponemella muris San Giorgi, Patho- 
logica rivista, 14, 1922, 461; Borrelia 
muris Bergey et al.. Manual, 2nd ed., 
1925, 435; Spirillum minus var. morsus 
muris Ruys, Cent. f. Bakt., I Abt., Orig., 
103, 1927, 270; Spirillum minus var. 



?nuris Ruys, ibid.; Spironevia minor 
Ford, Textb. of Bact., 1927, 962; Spiro- 
nema laverani Ford, ibid., 963 ; Spironema 
muris var. virginiana Ford, ibid., 963; 
Spirella morsusmuris Noguchi, in Jordan 
and P'alk, Newer Knowledge Bact. and 
Immun., 1928, 497; Spirella muris 
Noguchi, ibid.) From Latin, minus, 
less. 

Description taken from Adachi, Jour. 
E.xp. Med., 33, 1921, 647 and Giesberger, 
Inaug. Diss., Delft, 1936, 67. 

Short thick cells : 0.5 by 3.0 microns, 
having 2 or 3 windings which are thick, 
regular and spiral. Actively motile by 
means of bipolar tufts of flagella. Gram- 
negative. 

Has not been cultivated on artificial 
media. 

Aerobic, facultative. 

Pathogenic for man, monkej^s, rats, 
mice and guinea pigs. 

This species is regarded by some as a 
spirochaete. Because of its habitat and 
wide distribution it has been described 
under many different names. It is 
possible that some of these names indi- 
cate varieties or even separate species. 
See Beeson (Jour. Amer. Med. Assoc, 
123, 1943, 332) for important literature. 

Source : Found in the blood of rats and 
mice. 

Habitat : The cause of rat-bite fever. 
Widely distributed. 

7. Spirillum kutscheri Migula. {Spi- 
rillum undula majus Kutscher, Cent. f. 
Bakt., I Abt., 18, 1895, 614; Migula, 
Syst. d. Bakt., 2, 1900, 1024.) Named 
for Kutscher, the German bacteriologist 
who first isolated the organism. 

Stout threads, 1.5 microns in diameter. 
Wave lengths 10.5 to 12.5 microns. 
Width of spiral, 3 to 4.5 microns. May 
lose their spiral form on continued 
cultivation. Motile with tufts of flagella 
at the poles. Gram-negative. 

Gelatin colonies: Transparent, round, 
surface colonies. Deep colonies, dark 
brown. 



216 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Gelatin stab : Slow liquefaction. 

Agar colonies grow poorly, granular. 
Deep colonies yellowish-green to dark 
brown . 

Agar slant : Delicate, transparent 
growth. 

Potato: Limited growth. 

Volutin present. 

Catalase positive. 

Utilizes malic and succinic acids. 

Grows well on peptone broth. Also 
utilizes ammonia compounds. 

Optimum temperature, 22° to 27 °C. 

Source : Isolated from putrid materials 
and liquid manure. 

Habitat : Putrefying liquids. 

8. Spirillum volutans Ehrenberg. 
(Prototype, Vibrio spirillum Miiller, 
Animalcula infusoria, 1786; Ehrenberg, 
Die Infusionstierchen als Volkommene 
Organismen, 1838.) From M. L. volutin. 

Spirals 1 .5 microns in diameter. Wave 
length, 13 to 15 microns, width of spiral, 
4 to 5 microns. The largest of the spirilla. 
Slightly attenuated ends. Motile, pos- 
sessing a tuft of ten to fifteen flagella 
at each pole. Dark granules of volutin 
in the cytoplasm. Gram-negative. 

Migula (Syst. d. Bakt., 2, 1900, 1025) 
reports that this species has not been 
cultivated on artificial media, and that 
the cultures so described by Kutscher 
(Ztschr. f. Hyg., 20, 1895, 58) are of a 
different species which Migula names 
Spirillum giganteum. Vahle (Cent. f. 
Bakt., II Abt., 25, 1910, 237) later 
describes the cultural characters of an 
organism which he regards as identical 
with Kutscher 's organism. Giesberger 
(Inaug. Diss., Delft, 1936, 65) saw what 
he felt was the true Spirillum volutans 
but could not cultivate it. 

Optimum temperature 35°C. 

Habitat: Stagnant water. 



9. Spirillum lipoferum Beijerinck. 
(Azotobacter spirillum Beijerinck, Kon. 
Akad. Wetensch. Amsterdam, 30, 1923, 
431 quoted from Giesberger, Inaug. 
Diss., Delft, 1936, 24; Spirillum lipo- 
ferum Beijerinck, Cent. f. Bakt., II 
Abt., 63, 1925, 353; Chromatium lipo- 
ferum Bergey et al., Manual, 3rd ed., 
1930, 531.) From Greek, lipos, fat; 
Latin, fero, to bear. 

Curved cells with one-half to one spiral 
turn, containing minute fat droplets. 
These may deform the cells. Motile 
with lophotrichous flagella. Gram- 
negative. 

Calcium malate agar colonies: Circu- 
lar, small, transparent, dry. The malate 
is oxidized to calcium carbonate. Cells 
contain fat drops. 

Peptone agar colonies : More abundant 
development. Cells lack fat drops and 
are typically spirillum in form. 

Glucose peptone broth : Cells actively 
motile with large fat drops. 

Fixes atmospheric nitrogen in partially 
pure cultures, i.e., free from Azotobacter 
and Clostridium (Beijerinck, loc. cit.). 
Schroder (Cent. f. Bakt., II Abt., 85, 
1932, 17) failed to find fixation of nitrogen 
when she used cultures derived from a 
single cell. 

Aerobic. 

Optimum temperature 22°C. 

Beijerinck regards this as a transitional 
form between Spirillum and Azotobacter. 
Giesberger {loc. cit., p. 64-65) thinks it a 
Vibrio . 

Habitat: Garden soil. 

Appendix:* The following additional 
species have been mentioned in the 
literature. Many are inadequately de- 
scribed. Some may not belong here. 



* Prepared by Mr. Wm. C. Haynes, New York State Experiment Station, Geneva, 
New York, Jan., 1939; Revised by Capt. Wm. C. Haynes, Sn. C, Fort Bliss, Texas, 
.July, 1943. 



FA^nLY PSEUDOMOXADACEAE 



217 



Spirella canis Duboscq and Lebailly. 
(Compt. rend. Acad. Sci. Paris, 15Jt, 
1912, 835.) From the stomach of a dog. 

Spirillum amyliferum Van Tieghem. 
(Bull. Soc. botan. de France, 26, 1879, 
65.) Said to produce spores. Ford 
(Textb. of Bact., 1927, 364) thinks this 
organism was probably a spirochaete 
because of its mode of division. Found 
in frog spawn fungus of sugar factories. 

Spirillum attenuatum Warming. (Om 
nogle ved Danmarks Kyster levende 
Bakterier. Kjobenhavn, 1876; Spiro- 
soma attenuatum Migula, Syst. d. Bakt., 
2, 1900, 959.) Ford {loc. cit., 363) states 
that this incompletely described organ- 
ism would now be regarded either as a 
spirillum or as a spirochaete. From sea 
coast of Denmark. 

Spirillum cardiopyrogenes Sardjito. 
(Geneesk. Tijdschr. voor Xed. -Indie, 
72, 1932, 1359; ibid., 73, 1933, 822.) From 
lilood of a patient with pericarditis. 

Spirillum colossus Errera. (Rec. trav. 
bot. Bruxelle, 5, 1902; Abst. in Cent. f. 
Bakt., II Abt., 9, 1902, 608.) A giant 
form isolated from brackish sea water. 
Probably the same as Spirillum volutans 
Ehrenberg. 

Spirillum concentricum Kitasato. 
(Cent. f. Bakt., 3, 1888, 73.) Found in 
putrefying blood. 

Spirillum crassum Veillon and Repaci. 
(Ann. Inst. Past., 26, 1912, 300.) De- 
scribed as having peritrichous flagella. 
From lung lesions in human tuberculosis. 

Spirillum endo par agog icum Sorokin. 
(Cent. f. Bakt., 1, 1887, 465.) Described 
as producing spores in old cultures. 
From rain water in bark of poplar tree. 

Spirillum giganteum ^ligula. {Spiril- 
lum volutans Kutscher, Ztschr. f. Hyg., 
20, 1895, 58; Migula, Syst. d. Bakt., 2, 
1900, 1025.) From putrefying liquids. 

Spirillum hachaizae Kowalski. (Cent, 
f . Bakt., 16, 1894, 324 ; Spirillum hachaizi- 
cum Kowalski, ibid., 324; Spirochaete 
hachaizae Castellani and Chalmers, Man. 
Trop. Med., 1st ed., 1910, 316 ; Treponema 



hachaizae Brumpt, Nouveau Traite de 
Medecine, Paris, 4, 1922, 495.) Found 
in feces of cholera patients and also of 
healthy individuals. 

Spirillimi kolkwitzii Vislouch. (Jour, 
de Microbiol. (Russian), 1, 1914, 50.) 

Spirillum leiicomelaenum Perty. (Zur 
Kenntniss kleinster Lebensformen. 
Berne, 1852. Also see Koch, Mitt. 
Kais. Gesundheitsamte, /, 1881, 48.) 
From stagnant water. 

Spirilhun monospora Dobell. (Quart. 
Jour. Micr. Sci., 52, 1908, 121.) De- 
scribed as producing spores. From 
large intestine of frogs and toads. 

Spirillum nigrum Rist. (These med., 
Paris, 1898; see Cent. f. Bakt., I Abt., 
30, 1901, 299.) Strict anaerobe from pus. 

Spirillum ostreae Xoguchi. (Jour. 
Exp. Med., 3.^, 1921,295.) From oysters. 

Spirillum, periplaneticum Kunstler and 
Gineste. (Compt. rend. Soc. Biol. 
Paris, 61, 1906, 135.) From the intestine 
of the cockroach, Periplaneta amcricana. 

Spirillum pyogenes Mezincescu. 
(Cent. f. Bakt., I Abt., Orig., 35, 1904, 
201 ; Spirochaeta pyogenes Blanchard, 
Semaine Med., 26, 1906, 1; Treponema 
pyogenes Brumpt, Nouveau Traite de 
Medecine, Paris, 4, 1922, 511.) From a 
case of pyelitis calculosa. 

Spirillum, rappini De Toni and Trevi- 
san. (Spirochaete, Rappin, Contr. a 
I'Etude d. Bacter. de la Bouche a I'Etat 
normal, 1881, 68; De Toni and Trevisan, 
in Saccardo, Sylloge Fungorum, 8, 1889, 
1009.) From the stomach of a dog. 

Spirillum recti physeteris Beauregard. 
(Compt. rend. Acad. Sci. Paris, 125, 
1897,255.) From ambergris. 

Spirillum rugula (Miiller) Winter. 
{Vibrio rugula Miiller, Animalcula infu- 
soria, 1786; Cohn, Beitrage z. Biol. d. 
Pflanz., 1, Heft 2, 1872, 178; Bonhoff, 
Arch. f. Hyg., 26, 1896, 162; Winter, Die 
Pilze, in Rabenhorst's Kryptogamen- 
Flora, 1884.) Prazmowski found spores, 
but it is not certain his cultures were 
pure. Bonhoff also observed spores, but 



218 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



concluded that they were due to con- 
taminating organisms (Ford, Textb. of 
Bact., 1927, 360). From water. 

Spirillum sporiferum Migula. (Syst. 
d. Bakt., 2, 1900, 1028.) Produces 
spores. The spirals in which the spore 
formation is beginning are like Spirillum 
leucomelaenum Perty (Ford, loc. cit., 
336). Giesberger (loc. cit., p. 60) places 
this and other so-called spore-forming 
spirilla in Sporospirillum Orla-Jensen 
(Cent. f. Bakt., II Abt., 23, 1909, 340). 
From a bean infusion. 

Spirillum sputigenum Miller. (Die 
Mikroorganismen der Mundhohle. Leip- 
zig, 1892; Deutsche med. Wchnschr., 33, 
1906, 1 and 348. ) Hoffman and Prowazek 
(Cent. f. Bakt., I Abt., Orig., 4i, 1906, 
741) claim that Spirillum sputigenum 
hasperitrichousflagella. Giesberger {loc. 
cit., 63) places this in Selenomonas 
Prowazek (Cent. f. Bakt., I Abt., Orig., 
70, 1913, 36). Muhlens (Cent. f. Bakt., 
I Abt., Orig., 48, 1909, 525) reports 1 to 
3 flagella, the majority of the organisms 
having apparently a single thick fiagellum 
(a bunch of flagella) on the concave side 
(Ford, Zoc. cz7., 367). Anaerobic. From 
the buccal cavity. 

Spirillum slomachi Lehmann and Neu- 
mann. {Spirillum Form a, 13, y, 5 
Salomon, Cent. f. Bakt., I Abt., 19, 1896, 
433; Lehmann and Neumann, Bakt. 
Diag., 2 Aufl., 2, 1899, 362.) Found in 
stomach of dog, cat and rat. 

Paraspirillum vejdovskiiDoheW. (Arch, 
f. Protistenk., 24, 1911, 97.) Found only 



once in fresh water containing Oscilla- 
toria. Flagellate flexible spiral cells 
described as possessing a nucleus. This 
may be a protozoan. 

SpirohaciUus gigas Certes. (Bull. Soc. 
Zool. France, 14, 1889, 322; abst. in Ann. 
de Microgr., 2, 1889-1890, 137.) From 
water. 

Vibriothrix tonsillaris Tunnicliff and 
Jackson. (Organism from Actinomyces- 
like granules, Tunnicliff, Jour. Inf. Dis., 
38, 1926, 366; Tunnicliff and Jackson, 
ibid., 46, 1930, 12.) From tonsillar 
granules. May be identical with Lepto- 
thrix asteroide Mendel and as a Gram- 
negative, anaerobe may belong in Bac- 
teroides according to Rosebuiy (Bact. 
Rev., 8, 1944, 202). 

Vibriothrix zeylanica (Castellani) Cas- 
tellani. {Spirillum zeylanicum Castel- 
lani Jour. Ceylon Branch Brit. Med. 
Assoc, 7, 1910, 5 and Philipp. Jour. 
Sci., 5, No. 2, Sect. B., Medical Sciences, 
July, 1910; Vibrio zeylanicus Castellani, 
1913, Bacillus zeylanicus Castellani, 
1913 and Vibriothrix zeylanica Castellani, 
1917, quoted from Castellani and Chal- 
mers, Man. Trop. Med., 3rd ed., 1919, 
1069; Spirobacillus zeylanicus Castellani, 
Spagnuolo and Russo, Bull. Soc. Path. 
Exot., 11, 1918, 271.) Motile. Gram- 
negative. From cases of dysenteric 
enteritis in Ceylon. This is the type 
species of the genus Vibriothrix Castel- 
lani (see Castellani and Chalmers, loc. 
cit., 1068). 



FAMILY AZOTOBACTERIACEAE 



219 



FAMILY III. AZOTOBACTERIACEAE BERGEY, BREED AND MURRAY.* 

(Preprint, Manual, 5th ed., October, 1938, v and 71.) 

Cells without endospores. Relatively large rods or even cocci, sometimes almost 
yeast-like in appearance. The type of flagellation in this genus has been definitely 
established as peritrichous. Gram-negative. Obligate aerobes, usually growing in 
a film on the surface of the culture medium. Capable of fixing atmospheric nitrogen 
when provided with carbohydrate or other energ\^ source. Grow best on media 
deficient in nitrogen. Soil and water bacteria. 

There is a single genus. 

Genus I. Azotobacter Beijeririck. 

(Beijerinck, Cent. f. Bakt., II Abt., 7, 1901, 567; Azotomonas Orla-Jensen, Cent. f. 
Bakt., II Abt., 24, 1909, 444.) 

The definition is identical with that of the family. From Gr. azous, not living. 
French, azote, nitrogen; Gr. bakiron, rod, stick. 
The type species is Azotobacter chroococcu?7i Beijerinck. 



1. Azotobacter chroococcum Beijer- 
inck. (Cent. f. Bakt., II Abt., 7, 1901, 
567 and 9, 1902, 3; Bacillus azotobacter 
Lohnis and Hanzawa, Cent. f. Bakt., II 
Abt., 4^, 1914, 1; Bacillus chroococcus 
Buchanan, General Syst. Bact., Balti- 
more, 1925, 194.) From Gr. chroa, color; 
coccos, grain; M. L. sphere. 

According to Lohnis and Smith (Jour. 
Agr. Res., 23, 1923, 401) Azotobacter 
beijerinckii Lipman (New Jersey Agr. 
Exp. Sta. Rept., 25, 1904, 247), Azoto- 
bacter woodstownii Lipman (ibid.), Azoto- 
bacter smyrnii Lipman and Burgess (Cent, 
f. Bakt., II Abt., U, 1915, 504) and Azoto- 
bacter hilgardii Lipman (Science, 29, 
1909, 941) are identical with Azotobacter 
chroococcum. Greene (Soil Sci., 39, 
1935, 327) studied Azotobacter chroococ- 
cum and Azotobacter beijerinckii by 
chemical analyses and found the chemical 
composition of the cells to be practically 
identical, but different from that of 
Azotobacter vinelandii and Azotobacter 
agile. Smith (private communication) 
feels that Azotobacter beijerinckii is a 
non-pigmented rough strain of Azoto- 
bacter chroococcum. 



Grows in absence of organic nitrogen- 
Rods : 2.0 to 3.0 by 3.0 to 6.0 microns, 
occurring in pairs and packets and oc- 
casionally in chains. The cells show 
three or four refractile granules. The 
organisms are surrounded hj a slimy 
membrane of variable thickness, usually 
becoming brownish in older cultures, due 
possibly to the conversion of tyrosine to 
melanin. The coloring matter is insolu- 
ble in water, alcohol, ether and chloro- 
form. Motile bj' means of numerous 
peritrichous flagella (Hofer, Jour. Bact., 
47, 1944, 415.) Gram-negative. 

Gelatin colonies: Very small, circular, 
yellow, granular, later becoming yel- 
lowish-brown. 

Gelatin stab: Only slight growth in 
the stab. No liquefaction. 

Mannitol agar stab : Gray, may become 
brownish. 

Nutrient broth: No growth even in 
the presence of glucose ; peptone utilized 
with difficulty. 

Litmus milk : Becoming clearer in 10 
to 14 days. 
Potato: Glossy, barely visible, slimy 



* Revised by Dr. A. W. Hofer, New York State Experiment Station, Geneva, 
New York, June, 1938; further revision by Dr. A. W. Hofer, July, 1943. 



220 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



to wrinkled; may become yellowish, 
brownish-yellow or chocolate brown. 

The organism fixes atmospheric nitro- 
gen and gives off CO2, utilizing glucose 
and sucrose. Other generally used car- 
bon compounds are fructose, maltose, 
mannitol, inulin, dextrin, galactose, 
arabinose, starch, glycerol, ethyl alcohol, 
acetate, butyrate, citrate, lactate, mal- 
ate, propionate and succinate. 

Nitrate : Improves growth in amounts 
less than 1 gm. per liter ; greater amounts 
are toxic. 

Fixes nitrogen moderately actively. 

Chemical analysis : Four-day cultures 
grown upon mannitol agar (Greene, 
1935), when dried, are found to contain 
less than 0.5 per cent of hemicelluloses, 
less than 20 per cent of crude protein, 
less than 5 per cent of ash, and more 
than 30 per cent of lignin-like materials. 
The nitrogen fraction contains less than 
1 per cent of amide nitrogen, less than 1 
per cent of humin nitrogen and about 1 
per cent of basic nitrogen. 

Aerobic. 

Optimum temperature 25°C. to 28°C. 

Distinctive characters : Inability to 
grow in peptone media, even in the 
presence of glucose; frequent occur- 
rence of a dark brown or black pigment. 

Source : Isolated from soil. 

Habitat: Occurs naturally in the 
majority of neutral or alkaline field soils. 

2. Azotobacter agile Beijerinck. 
(Cent. f. Bakt., II Abt., 7, 1901,577.) 
From L. agilis, agile, quick. 

In studies on the chemical composition 
of cells Greene (Soil Sci., 39, 1935, 327) 
found Azotobacter vinelandii Lipman 
(New Jersey Agr. Exp. Sta. Rept., 24, 
1903, 238) to be very similar to Azoto- 
bacter agile Beijerinck. Smith and Loh- 
nis (Jour. Agr. Res., 23, 1923, 401) agree 
and state furthermore that the two are 
identical; they believe also that Azoto- 
bacter vitreiim Lohnis and Westermann 
(Cent. f. Bakt., II Abt., 22, 1908, 234) 
is another synonym of Azotobacter agile. 



Smith (private communication) states 
that Azotobacter vitreum is a very weak 
growing, smooth strain of Azotobacter 
agile. Kluyver and van Reenen (Arch. 
Mikrobiol., 4, 1933, 299) feel that a dis- 
tinction should be made between Azoto- 
bacter agile and Azotobacter vinelandii. 
In regard to the former, Kluyver and 
van den Bout (Arch. Mikrobiol., 7, 1936, 
263) suggest that it be further subdivided 
into Azotobacter agile and Azotobacter 
agile var. atypica, the latter referring to 
an Azotobacter agile form that fails to 
produce pigment. 

Rods: 4 to 6 microns in length, almost 
spherical. Actively motile by means of 
numerous peritrichous flagella (Hofer, 
loc. oil). Some strains are reported to 
be non-motile. Gram-negative. 

Grows in absence of organic nitrogen. 

Gelatin : No liquefaction. 

Mannitol agar colonies : Circular, gray- 
ish white, translucent with whitish 
center. 

Washed agar colonies : Show slight 
bluish-green fluorescence. 

Mannitol agar slant: Grayish, trans- 
lucent, fluorescent. 

Plain agar slant : Yellowish-white, 
smooth, glistening, translucent with 
opaque center. 

Broth: Turbid, with sediment. 

Litmus milk : Becoming clear in 10 to 
14 days. 

Potato : Yellowish-white, slimy, be- 
coming yellowish-brown. 

In the presence of organic acids, a 
greenish or reddish pigment is formed. 

The organism fixes atmospheric nitro- 
gen actively, and gives off CO2. 

Aerobic. 

Chemical analysis : Four-day cultures 
grown upon mannitol agar (Greene, 1935), 
when dried, contain more than 4 per cent 
of hemicelluloses, more than 45 per cent 
of crude protein, more than 7 per cent 
of ash, and less than 4 per cent of lignin- 
like materials. The nitrogen fraction 
contains more than 1 per cent amide 
nitrogen, more than 1 per cent humin 



FAMILY AZOTOBACTERIACEAE 



221 



nitrogen, and 2 per cent or more of basic 
nitrogen. 

Optimum temperature 25 °C to 28 °C. 

Distinctive characters : Lack of a 
brown pigment ; occasional fluorescence ; 
growth in peptone broth containing 
glucose. 

Source : Originally isolated from canal 
water at Delft. 

Habitat: Occurs in water and soil. 

3. Azotobacter indicum Starkey and 
De. (Soil Sci. 47, 337, 1939.) From L. 
indict/a, of India. 

Rods: Ellipsoidal, from U. 5 to 1.2 by 1.7 
to 2.7 microns when grown on nitrogen 
free glucose agar. One of the distinctive 
characteristics is the presence of two 
large, round, highlj'' refractive bodies in 
the cells, one usually at each end. Mo- 
tile by means of numerous peritrichous 
fiagella (Hofer. loc. cit.). Gram-nega- 
tive. 

The organism grows slowly but in 
time produces large amounts of slime. 



Has high acid tolerance, since it grows 
from pH 3 to 9. 

Sucrose or glucose agar plates : Colonies 
are colorless, round, very much raised, 
and uniformly turbid, having much the 
appearance of heavy starch paste. Af- 
ter two weeks, a buff to light brown color 
develop;. 

Mannitol agar slant : Grows very 
poorly. 

Peptone agar slant with 0.5 per cent 
glucose: Limited grayish growth. 

Nutrient broth : No growth. 

Liquid media generally : Turbidity 
with some sediment. 

Fixes atmospheric nitrogen readily 
with either glucose or sucrose as source 
of energy. 

Aerobic. 

Optimum temperature: 30°C. 

Distinctive characters : Tolerance of 
acidity, wide limits of pH tolerated, 
abundant slime production, large glob- 
ules of fat within cells. 

Source : Soils of India. 

Habitat : Soils. 



Appendix I : The relationship of the following species to the species placed in .420- 
tobacler is not yet entirely clear. 

Genus Azotomonas Stapp. 

(Cent. f. Bakt., II Abt., 102, 1940, 18; not Azotomonas Orla-Jen.sen, Cent. f. Bakt., 

II Abt., 24, 1909, 484.) 

Rod to coccus-shaped aerobic bacteria, motile by means of 1 to 3 polar Hagella. Xo 
endospores. No fat-like reserve food granules in the cells. Form acid and gas from 
glucose, and other sugars and alcohols. Form indole. Chemo-heterotrophic. Many 
carbon compounds other than sugars used as sources of energy. Active in the fixation 
of atmospheric nitrogen. Live in soil. From Gr. azvus, not living. French, azote, 
nitrogen; Greek, monas, a unit; M. L. monad. 

The type species is Azotomonas insolita. 

Azotomonas insolita Stapp. (Ab- microns. Motile with one to three polar 
stracts of Communications, Third In- flagella. Gram-negative. 



ternat. Congr. for Microbiol.. Sect. VIII, 
1939, 306; abst.in Proc. Soil Sci. Soc. of 
America, 4, 1939, 244; Cent. f. Bakt., II 
Abt., 102, 1940, 1.) From Latin insolilus, 
unusual . 



Gelatin: No liquefaction. 
Agar slant: Glistening white growth. 
Agar colonics: Flat, whitish, edge 
entire. Weakly fluorescent. 
Broth : Strong turbidity. Sediment. 



Coccoid rods: 0.6 to 1.2 bv 0.6 to 1.8 Pellicle. 



222 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Milk: No change. 

Potato : Growth somewhat dry, not 
slimy, dirty gray, spreading. 

Nitrites produced from nitrates. 

Fixes nitrogen. 

Ammonium salts utilized. 

Acid and gas from adonitol, arabinose, 
dextrin, glucose, galactose, glycerine, 
inositol, lactose, fructose, maltose, man- 
nitol, mannose, raffinose, rhamnose, sali- 
cin, sorbitol, starch, sucrose and xylose. 



Starch is hydrolyzed. 

Hydrogen sulfide produced. 

Optimum temperature 25° to 30 °C. 
Minimum 7° to 9.5°C. Maximum 48°C. 
Good growth at 37 °C. Thermal death 
point 60 °C. 

Limits of pH 3.3 to 9.5. 

Aerobic. 

Source : From a mixture of chopped 
cotton husks and rice hulls. 

Habitat: Soil. 



FAMILY- RmzaMACH^E-- 223 

FAMILY IV. RHIZOBL\CEAE COXX. 

(Jour. Bact., 36, 1938, 321.) 
Cells without endospores, rod-shaped, sparsely flagellated (one polar or lateral flagel- 
lum, or 2 to 4 peritrichous ones) ; some species non-motile. Usually Gram-negative. 
One genus (Chrotnobacterium) produces a violet pigment. Grow aerobically on ordi- 
nary culture media containing glucose. Glucose and sometimes other carbohydrates 
are utilized, without appreciable acid formation. Saprophytes, symbionts and 
pathogens. The latter are usually plant pathogens forming abnormal growths on 
roots and stems. 

Key to genera of family Rhizobiaceae. 

I. Cells capable of fixing free nitrogen when growing symbiotically on the roots of 
Leguminosae. 

Genus I. Rhizobium, p. 223. 
II. Either plant pathogens which attack roots or produce hypertrophies on stems ; or 
free-living non-chromogenic soil or water forms. Do not fix nitrogen. 

Genus II. Agrobacterium, p. 227. 
III. Usually free-living soil and water forms which produce a violet chromogenesis. 

Genus III. Chroniobacterium, p. 231. 

Genus I. Rhizobium Frank.* 

{Phytomyxa Schroeter, in Cohn, Kryptogamen-Flora von Schlesien, S, 1886, 134; 
Frank, Ber. d. deut. bot. Gesellsch., 7, 1889, 380; Rhizobacterium Kirchner, Beitr. z. 
Biol. d. Pflanzen, 7, 1895, 221; Rhizomonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 
1909, 328.) From Greek rhiza, root; bios, life. 

Rods: 0.5-0.9 by 1.2-3.0 microns. Motile when young, commonly changing to bac- 
teroidal forms (a) upon artificial culture media containing alkaloids or glucosides, or 
in which acidity is increased; or (b) during symbiosis within the nodule. Gram- 
negative. Aerobic, heterotrophic, growing best with extracts of yeast, malt or other 
plant materials. Xitrates may be reduced to nitrites. Nitrites are not utilized. 
Gelatin is not liquefied or is very slightly liquefied after long incubation. Optimum 
temperature 25°C. This group is capable of producing nodules on the roots of 
Leguminosae, and of fixing free nitrogen during this symbiosis. 

The type species is Rhizobium leguminosarum Frank. 

Key to the species of genus Rhizobium. 
1. Litmus milk alkaline. 

a. Formation of serum zone in milk. 

b. Moderate growth, slight acid reaction on yeast water agar plus mono-, di- 
and trisaccharides. 

c. Causes formation of root nodules on species of the genera Lathyrus, 
Pisum, Vicia and Lens. Bacteroids irregular with x, y, star-, and 
club-shaped forms; rods peritrichous when young. 

1. Rhizobium leguminosarum. 

cc. Causes formation of root nodules on Phaseolus vulgaris, P. multiflorus 
and P. angustif alius . Bacteroids vacuolated rods, few branched 
forms ; j'oung cells peritrichous. 

2. Rhizobium phaseoli. 

* The genus Rhizobium was revised by Dr. and Mrs. O. N. Allen under the direction 
of Prof. E. B. Fred and Prof. I. L. Baldwin, Univ. of Wisconsin, Madison, Wis., Jan., 
1938; further revision by Dr. O. X. Allen, Jan., 1943. 



224 MANUAL OF DETERMINATIVE BACTERIOLOGY 

ccc. Causes formation of nodules on species of genus Trifolium. Bac- 
teroids pear-sliaped, swollen, vacuolated. Pentoses usually not 
fermented. 

3. Rhizobium trijolii. 
aa. No serum zone formed in milk. 

b. Scant growth, alkaline reaction on yeast water agar plus most carbohydrates. 
c. Causes formation of nodules on species of genus Lupinus and on Orni- 
thopus sativus. Bacteroids vacuolated, rods seldom branched. 

4. Rhizohium lupini. 

cc. Causes formation of nodules on Soja max. Bacteroids long slender 
rods, seldom vacuolated or branched; young cells monotrichous. 

5. Rhizobium japonicum.* 
2. Litmus milk acid. 

a. Formation of serum zone in milk. 

b. Moderate growth, slight acid reaction on yeast w^ater agar plus mono-, di- 
and trisaccharides. 

c. Causes formation of root nodules on species of the genera Melilotus, 
Medicago, and Trigonella. Bacteroids club-shaped, branched, young 



cells peritrichous. 



6. Rhizobium meliloti. 



1. Rhizobium leguminosarum Frank Naturreiche. 2 Theil, Botanik, III Abt., 

emend. Baldwin and Fred. (Frank, Kryptogamen, Sec. 914, 1877, 1944; 

Landwirtschaftliche Jahrbiicher, 79, 1890, Schinzia leguminosarum Frank (all spe- 

563; Rhizobium polymorphum Dangeard, cies), Bot. Ztg., 37, 1879, 377; Phyto- 

Rhizobiumfabae'Dangeard,'LeBotsimste, viyxa leguminosarum Schroeter (all ex- 

Ser. 16, 1926, 192-194 ; Baldwin and Fred, cept Rhizobium lupini), in Cohn, Krypto- 

Jour. Bact., 17, 1929, 146.) From Latin, gamen-Flora von Schlesien, 3, I, 1886, 

of the legume family (Leguminosae) . 135; Bacillus radicicola Beijerinck (all 

species), Bot. Ztg., 46, 1888, 726; Bacillus 

Note: The following binomials have fabae Beijerinck (from broad bean) and 

been used for species of this genus. The Bacillus oi-nilhopi Be'iierinck (from ser- 

names given were used by their authors radella), Bot. Ztg., 48, 1890, 837; Clado- 

to cover one or more of the species here chytriuin tuberculorum Vuillemin (all 

recognized as belonging to the genus species?), Ann. Sci. Agron. Franc, et 

Rhizobium. Where a question mark (?) Etrang., 5, I, 1888, 193; Bacterium 

is used it indicates that the species was radicicola Prazmowski (all species), 

too poorly described to be recognizable Landw. Vers. Sta., 37, 1890, 204; Rhizo- 

today. Schinzia cellulicola Frank, 1877 femmnmiobiZe Schneider (several species) 

(all species) Leunis, Synopsis der drei Rhizobium curvum Schneider (?), Rhizo- 

* No specific name has been proposed for the organism causing the formation of 
nodules on plants that are members of the so-called "cowpea" group. Data showing 
possible inter-relationships of certain plant species of the soybean and cowpea cross- 
inoculation groups prompted Walker and Brown (Soil Science, 39, 1935, 221-225) to 
propose a consolidation of the two groups to be recognized as being inoculated by a 
single species, Rhizobium japonicum. Results obtained recently by Reid and Bald- 
win (Proc. Soil Sci. Soc. Amer. for 1936, 1, 1937, 219) show these inter-relationships to 
include the lupine group also. 



FAMILY HHIZOBIACEAE 



225 



hium frankii var. majus and var. mi- 
nus Schneider (?), Rhizohium nodosum 
Schneider (?), Rhizohium dubium 
Schneider (?), Bui. Torrej' Bot. Club, 
19, 1892, 213; Rhizohium sphaeroides 
Schneider (?), Ber. deut. bot. Gesell., 12, 
1894, 16; Bacillus tuberigenus Gonner- 
mann and Micrococcus tuberigenus Gon- 
nermann, Landw. Jahrb., 33, 1894, 654, 
657, are thought bj" Fred, Baldwin and 
McCoy (University of Wisconsin, Stud- 
ies in Science, Xo. 5, 1932, 140) not to be 
true noduJe organisms and to be too 
poorly described to be recognizable to- 
day ; Rhyzobium pasteurianum Maze (all 
species), Ann. Inst. Pasteur, 13, 1899, 
146; Pseudorhizobium ramosnm Hartleb 
(?) (Chem. Zeit., U, 1900, 887) (used 
for noninfective culture claimed b}" 
Stutzer (Mitt. Landw. Inst. Breslau, /, 
Heft 3, 1900, 63) to be }>;enuine root nod- 
ule organism) ; Rhizohium radicicola 
Hiltner and Stormer (several species) 
and Rhizohium heijerinckii Hiltner and 
Stormer (from lupine, serradella and soy 
bean), Arb. Biol. Abt. f. Land-u. Forst- 
wirthschaft a. K. Gesundheitsamte, 3, 
1903, 269; Pseudomonas radicicola ]\Ioore 
(all species), U. S. Dept. Agr. Bur. 
Plant Ind., Bui. 71, 1905, 27; Rhizomonas 
heijerinckii Orla-Jensen and Rhizomonas 
radicicola Orla-Jensen (see Hiltner and 
Stormer), Cent. f. Bakt., II Abt., 22, 
1909, 328; Bacillus or Bacterium radici- 
cola Lohnis and Hansen (peritrichous 
species). Jour. Agr. Research, 20, 1921, 
554 ; Rhizohium radicicolum Bergej' et al. , 
Manual, 1st ed., 1923, 40 (monotrichous 
species) ; Rhizohium loti Dangeard (from 
lotus), Rhizohium simplex Dangeard 
(from sainfoin), Rhizohium torulosum 
Dangeard (from Scotch broom), Le 
Botaniste, Ser. 16, 1926, 195-197. 

Rods: 0.5 to 0.9 by 1.2 to 3.0 microns. 
Motile with peritrichous flagella. Bac- 
teroids commonly irregular with x, y, 
star- and club-shaped forms. Vacuolate 
forms predominate. Gram-negative. 

Growth on mannitol agar is rapid, with 
tendency to spread. Streak is raised, 



glistening, semi-translucent, white, slimy 
and occasionally viscous. Considerable 
gum is formed. 

Slight acid production from glucose, 
galactose, mannose, lactose and maltose. 

Aerobic. 

Optimum temperature 25°C. 

Source : Root nodules on Lathyrus, 
Pisum (poa), Vicia (vetch) and Lens 
(lentil). 

Habitat : Widely distributed in soils 
where the above mentioned legumes are 
grown 

2. Rhizobiumphaseoli Dangeard. (Le 
Botaniste, Ser. 16, 1926, 197.) From 
Latin, phascolus, bean; M. L. Phaseolus, 
a generic name. 

Rods : Motile with peritrichous fla- 
gella. Bacteroids are usuallj^ rod-shaped, 
often vacuolated with few branched 
forms. Usually smaller than in Rhizo- 
hium leguminosariim and R. trifolii. 
Gram -negative. 

Growth on mannitol agar is rapid with 
tendency to spread. Streak inoculation 
is raised, glistening, semi-translucent, 
white, slimJ^ Occasionally mucilagi- 
nous but this character is not so marked 
as in Rhizohium trifolii. 

Very slight acid formation from glu- 
cose, galactose, mannose, sucrose and 
lactose. 

Aerobic. 

Optimum temperature 25 ''C. 

Source : Root nodules of Phaseolus 
vulgaris (kidney bean), P. ayigustifolius 
(bean) and P. multiflorus (scarlet run- 
ner). (Burrill and Hansen, 111. Agr. 
Exp. Sta. Bui. 202, 1917, 137.) 

Habitat : Widely distributed in the 
soils in which beans are grown. 

3. Rhizobium trifolii Dangeard. (Le 
Botaniste, Ser. 16, 1926, 191.) From 
M. L. Trifolium, a generic name. 

Rods : Motile with peritrichous fla- 
gella. Bacteroids from nodules are pear- 
shaped, swollen and vacuolated. Rarely 
X and y shapes. Gram-negative. 



226 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Growth on mannitol agar is rapid. Tlic 
colonies are white becoming turbid with 
age. Frequently mucilaginous. Streak 
cultures transparent at first. Growth 
mucilaginous later flowing down the 
agar slant and accumulating as a slimy 
mass at the bottom. Produces large 
amounts of gum. 

Slight acid production from glucose, 
galactose, mannose, lactose and maltose. 

Aerobic. 

Optimum temperature 25°C. 

Source : Root nodules of species of 
Trifolium (clover). 

Habitat: Widely distributed in the 
soils where clover grows. 

4. Rhizobixim lupini (Schroeter) Eck- 
hardt, Baldwin and Fred. {Phytomyxa 
lupini Schroeter, in Cohn, Kryptogamen- 
Flora von Schlesien, 3, I, 1886, 135; 
Rhizobium minimum Dangeard, Le Bo- 
taniste, Ser. 16, 1926, 198; Eckhardt, 
Baldwin and Fred, Jour. Bact., 21, 1931, 
273.) From Latin, Lupinus, lupine. 

Rods : Motile with flagella 1 to 4, usually 
2 or 3. Bacteroids are vacuolate rods, 
seldom if ever branched. Gram-negative. 

Growth on yeast water, mannitol agar 
is scant to moderate with alkaline 
reaction. 

Beef -peptone gelatin : Little growth 
with extremely slow liquefaction. 

On galactose an alkaline reaction serves 
to differentiate Rhizobium lupini from 
all fast-growing rhizobia (R. phaseoli, 
R. meliloti, R. trifolii, and R. legumino- 
sarum). An initial alkaline reaction fol- 
lowed more quickly by an acid reaction 
on rhamnose and xylose separates R. 
lupini from slow-growing R. japonicxim 
and the Rhizobium sp. from cow pea. 

In general Rhizobium lupini produces 
slight to moderate acidity on pentose 
sugars and no change or alkaline reaction 
on hexoses, disaccharides and trisac- 
charides. 

Litmus milk : No serum zone, no re- 
duction, and a slight alkaline reaction. 



Meager growth on potato and parsnip 
slants, and carrot agar. 

Aerobic. 

Optimum temperature 25°C. 

Source : Root nodules on Lupinus 
(lupine), Serradella and Ornithopus. 

Habitat : Widely distributed in soils 
in which these legumes grow. 

5. Rhizobium japonicum (Kirchner) 
Buchanan. (Rhizobacterium japonicum 
Kirchner, Beitrage zur Biol. d. Pflanzen, 
7, 1895, 213; Pseudomonas japonica 
Lohnis and Hansen, Bacterium japoni- 
cum Lohnis and Hansen, Jour. Agr. 
Res., 20, 1921, 551; Rhizobium sojae 
Dangeard, Le Botaniste, Ser. 16, 1926, 
200; Buchanan, Proc. Iowa Acad. Sci., 
33, 1926, 81.) From M. L., of Japan. 

Rods : Motile with monotrichous fla- 
gella. Bacteroids of nodules are long 
and slender with only occasional branched 
and swollen forms. Gram-negative. 

Growth on mannitol agar is slow and 
scant. The streak is slightly raised, 
glistening, opaque, white, butyrous, with 
little gum formation. 

Pentose sugars give better growth than 
the hexoses. 

Little if any acid formed from carbo- 
hydrates. Acid slowly formed from 
xylose and arabinose. 

Aerobic. 

Optimum temperature 25°C. 

Source : Root nodules on Soja max 
(soy bean). 

Habitat : Widely distributed in soils 
where soy beans are grown. 

6. Rhizobium meliloti Dangeard. (Le 
Botaniste, Ser. 16, 1926, 194.) From 
Greek, melilot, a kind of clover; M. L., 
Melilotus. 

Rods: Motile with peritrichous fla- 
gella. Bacteroids club-shaped and 
branched. Gram-negative. 

Growth on mannitol agar is fairly rapid. 
The streak is raised, glistening, opaque, 
pearly white, butyrous. Considerable 
gum is formed. 



FAMILY RHIZOBIACEAE 227 

Acid from glucose, galactose, mannose Note : See Monograph on Root Nodule 

and sucrose. Bacteria and Leguminous Plants by E. 

Aerobic. B. Fred, I. L. Baldwin and Elizabeth 

Optimum temperature 2o°C. McCoy, University of Wisconsin Studies 

Source : Root nodules of Melilotus in Science, Madison, No. 5, 1932, xx + 

(sweet clover), Medicago, and Trigonella. 343 pp. for a more complete discussion of 

Habitat : Widelj' distributed in soils this group with an extensive bibliogra- 

in which these legumes grow. phy. 

Genus II. Agrobacterium Conn.* 

(.Jour. Bact., 44, 1942, 359.) From Greek, agrus, a field; M.L., bacterium, a small 

rod. 

Small, short rods which are typically motile with 1 to 4 peritrichous flagella (if 
only one flagellum, lateral attachment is as common as polar). Ordinarily Gram- 
negative. On ordinary culture media, they do not produce visible gas nor sufficient 
acid to be detectable by litmus. In synthetic media, enough CO2 may be produced to 
show acid with brom thymol blue, or sometimes with brom cresol purple. Gelatin 
is either very slowly liquefied or not at all. Free nitrogen cannot be fixed ; but other 
inorganic forms of nitrogen (nitrates or ammonium salts) can ordinarilj^ be utilized. 
Optimum temperature, 25° to 30°C. Habitat: Soil, or plant roots in the soil; or the 
stems of plants where they produce hypertrophies. 

The type species is Agrobacteriitm tumefaciens (Smith and Townsend) Conn. 

Key to the species of genus Agrobacterium. 

I. Plant pathogens. Produce browning of mannitol-calcium-glycerophosphate agar. 
Nitrate reduction weak or none. 

A. Nitrite produced from nitrate toaslight extent. Galls produced on plant roots. 

1. Agrobacterium tumefaciens. 

B. Nitrite not produced from nitrate. 

1. Pathogenic to apples. 

2. Agrobacteriitm rhizogenes. 

2. Pathogenic to raspberries and blackberries. 

3. Agrobacterium rubi. 

II. Not pathogenic to plants. Produces browning in mannitol-calcium-glycerophos- 
phate agar. Nitrate reduction vigorous, with disappearance of the nitrate. 

4. Agrobacterium radiobacier. 

1. Agrobacterium tumefaciens (Smith 1923, 189; Conn, Jour. Bact., 44, 1942, 

and Town.send) Conn. (Bacterium tume- 359.) From Latin tumefaciens, swell- 

faciens Erw. Smith and Townsend, Sci- ing up, producing a tumor, 

ence, N. S. 25, 1907, 672; Pseudomonas Probable sj'nonyms : Bacillus ampelop- 

tumefaciens Stevens, The Fungi which sorae Trevisan, in Saccardo, S3dloge 

Cause Plant Disease, 1913, 35; Bacillus Fungorum, 8, 1889, 983; Bacillus am- 

tumefaciens Holland, Jour. Bact., 5, 1920, pelopsorae Trevisan emend. Cavara, Staz. 

220; not Bacillus tumefaciens Wilson, Sperim. Agara. Ital. Modena, 30, 1897, 

Lancet, 1, 1919, 675; Phytomonas tume- 483; see Elliott, Bact. Plant Pathogens, 

faciens Bergey et al.. Manual, 1st ed., 1930, 235. 

* Prepared by Prof. H. J. Conn, New York State Experiment Station, Geneva, 
New York, September, 1943. 



228 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Among the synonyms listed in previous 
editions of the Manual has been Poly- 
monas tumefaciens Lieske, Cent. f. 
Bakt., I Abt., Orig., 108, 1928, 118. This 
is only a partial synonym, however, as 
its author described it as the cause of 
animal and human cancer, of which he 
regarded crown-gall of plants as merely 
a phase : for the origin of this theory, see 
Smith and Townsend, Sci., N.S. 25, 1907, 
671, and Smith, Jour. Cancer Res., 7, 
1922, 1-105. 

Description taken from the following: 
Riker, Banfield, Wright, Keitt and 
Sagen, Jour. Agr. Res., 41, 1930, 507; 
Sagen, Riker and Baldwin, Jour.Bact., 
£8, 1934, 571; Hendrickson, Baldwin and 
Riker, Jour. Bact., 28, 1934, 597. 

Rods: 0.7 to 0.8 by 2.5 to 3.0 microns, 
occurring singly or in pairs. Capsules. 
Motile with 1 to 4 flagella. Gram- 
negative. 

Agar colonies: Small, white, circular, 
smooth, glistening, translucent, entire. 

Broth: Slightly turbid, with thin pel- 
licle. 

Litmus milk: Slow coagulation. Lit- 
mus reduced. Neutral to alkaline. 

Nitrites produced from nitrates to a 
very slight extent. 

Indole: Slight amount. 

Slight acid from glucose, fructose, 
arabinose, galactose, mannitol and salicin. 

Starch not hydrolyzed. 

Optimum temperature 25° to 28°C. 

Facultative anaerobe. 

Distinctive characters : Causes a gall 
formation parenchymatous in character 
which because of its soft nature is sub- 
ject to injury and decay. 

Agrobacterium tumefaciens strongly ab- 
sorbs Congo red and aniline blue in con- 
trast to little or no absorption by A. 
rhizogenes. A. tumefaciens makes abun- 
dant growth on sodium selenite agar and 
calcium glycerophosphate medium with 
mannitol in contrast to no growth or a 
very slight trace by A. rhizogenes (Hen- 
drickson et al., Jour. Bact., 28, 1934, 
597). 



Source: Isolated from galls on plants. 

Habitat : Causes galls on Paris daisy 

and cross-inoculable on over 40 families. 

2. Agrobacterium rhizogenes (Riker 
et al.) Conn. (.Bacterium rhizogenes 
Riker, Banfield, Wright, Keitt and Sagen, 
Jour. Agr. Res., 41, 1930, 536; Phyto- 
monas rhizogenes Riker et al., ibid., 
536; Pseudomonas rhizogenes Riker et 
al., ibid. 536; Conn, Jour. Bact., 44, 1942, 
359.) From Greek, rhiza, root; genes, 
producing. 

Rods: 0.4 by 1.4 microns, occurring 
singly. Motile with one to 4 flagella. 
Encapsulated. Not acid-fast. Gram- 
negative. 

Gelatin : No liquefaction. 

Agar colonies : Circular, smooth, con- 
ve-x, finely granular; optical characters, 
translucent through gray to almost white. 

Agar slant: Moderate, filiform, trans- 
lucent, raised, smooth, slimy. 

Broth: Turbid, with heavy pellicle. 

Litmus milk: Acid, slow reduction. 

Indole not formed. 

Nitrites not produced from nitrates. 

Acid but not gas from arabinose, xy- 
lose, rhamnose, glucose, galactose, man- 
nose, maltose, lactose, salicin and ery- 
thritol. No acid or gas from fructose, 
sucrose, raffinose, melezitose, starch, 
dextrin, inulin, aesculin, dulcitol or man- 
nitol. 

Starch not hydrolyzed. 

Optimum temperature 20° to 28°C. 

Aerobic. 

Distinctive characters : Agrobacterium 
rhizogenes differs from Agrobacterium 
tumefaciens by stimulating root forma- 
tion instead of soft parenchymatous 
crown galls. A. rhizogenes lacks ability 
of A. tumefaciens to utilize simple 
nitrogenous compounds as KNO3. A. 
rhizogenes absorbs congo red and brom 
thymol blue slightly and aniline blue not 
at all. Will not grow on sodium selenite 
agar (see A. tumefaciens for response 
to same materials). Does not infect 
tomato . 



FAMILY RHIZOBIACEAE 



229 



Sources : Description made from ten 
cultures isolated from hairy -root of apple 
and other plants. 

Habitat: Pathogenic on apple, etc. 

3. Agrobacterium rubi (Hildebrand) 
Starr and Weiss. {Phytomonas rubi 
Hildebrand, Jour. Agr. Res., 61, 1940, 
694; Bacterium rubi Hildebrand, ibid., 
694; Pseudomonas rubi Hildebrand, ibid., 
694; Banfield, Phytopath., 20, 1930, 123; 
Pinckard, Jour. Agr. Res., 50, 1935, 933; 
Starr and Weiss, Phytopath., 33, 1943, 
316.) From Latin, rubus, blackberrj' 
bush; M. L., Rubus, a generic name. 

Rods: 0.6 by 1.7 microns. Singly, in 
pairs or short chains. IMotile with 1 to 4 
flagella. Gram-negative. 

Gelatin : No liquefaction. 

Potato-mannitol-agar slants: Growth 
slow, moderate, filiform, white to creamy- 
white, with butyrous consistency later 
becoming leathery. 

Broth: Turbid in 36 to 48 hours. 

Milk: A slight serum-zone, pink color, 
acid and curd formed. 

Nitrites not produced from nitrates. 

Ferric ammonium citrate, uric acid, 
oxamide, succinimide, 1-asparagine, 1- 
tjTosine, 1-cystine, d-glutamic acid and 
yeast extract can be used as a source of 
nitrogen (Pinckard, loc. cit.). 

Hydrogen sulfide not formed. 

Indole not formed. 

Acid from glucose, d-galactose, d- 
mannose, d-fructose, d-xylose, d-arabin- 
ose, sucrose, and maltose. None from 
lactose (Pinckard, loc. cit.). 

Starch not hydrolyzed. 

Optimum temperature 28°C. Mini- 
mum 8°C. and maximum 36°C. (Pinck- 
ard, loc. cit.). 

Distinctive characters. Differs from 
Agrobacterium tumefaciens in that it 
does not utilize nitrates, and grows much 
more slowly on ordinary media. Infects 
only members of the genus Rubus. Starr 
and Weiss (Phytopath., 33, 1943, 317) 
state that this species unlike Agrobac- 
ferivm itimefaciens and Agrobacterium 



rhizogcncs does not utilize asparagin as a 
sole source of carbon and nitrogen. 

Source: Isolated by Banfield {loc. cit.) 
and by Hildebrand {loc. cit.) from rasp- 
berry canes, Rubus spp. 

Habitat : Pathogenic on black and 
purple cane raspberries, and blackberries, 
and to a lesser extent on red raspberries. 

4. Agrobacterium radiobacter (Bei- 
jerinck and van Delden) Conn. {Bacillus 
radiobacter Beijerinck and van Delden, 
Cent. f. Bakt., II Abt., 9, 1902, 3; Bac- 
terium radiobacter Lohnis, Cent, f . Bakt., 
II Abt., 14, 1905, 589; Rhizobium radio- 
bacter Pribram, Klassifikation der Schizo- 
myceten, Leipzig, 1933 ,5Z lAchromobacter 
radiobacter Bergey ct al., Manual, 4th ed., 
1934, 230; Alcaligenes radiobacter Conn, 
in Manual, 5th ed., 1939, 97; Conn, Jour. 
Bact., U, 1942,359.) From Latin, mdn^s, 
the spoke of a wheel; Latin, bactrum, 
a rod. 

Small rods, 0.15 to 0.75 l^y 0.3 to 2.3 
microns, occurring singly, in pairs and 
under certain conditions, in star-shaped 
clusters. Motile with one to four flagella. 
Prevailingly Gram-negative ; but an occa- 
sional culture is variable. 

Nutrient gelatin stab : No liquefaction. 

Agar slant: Flat, whitish slimy layer. 

Mannitol-calcium-glycerophosphate- 
agar streak plates: Abundant, raised, 
slimy growth surrounded by a brown 
halo with an outer zone of white precip- 
itate (Riker et al.. Jour. Agr. Res., 41, 
1930, 524). 

Broth: Turbid; with heavy ring or 
pellicle if veal infusion is present. 

Litmus milk : Serum zone with pellicle 
in one week; usually turns a chocolate 
brown in 2 weeks; same in plain milk, 
but with less browning. 

Potato : Raised slimy mass becoming 
brownish; potato may be browned. 

Nitrates disappear (assimilated or 
reduced). 

Starch not hydrolyzed. 

No organic acid or visible gas from 
sugars; nearly all sugars, glycerol and 



230 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



mannitol are utilized with the produc- 
tion of CO2. 

Optimum temperature 28°C. Mini- 
mum near 1°C. Maximum 45°C. 

Aerobic. 

Media containing KNO3, K2HPO4, and 
glycerol, ethyl or propyl alcohol become 
alkaline to phenol red. (Sagen, Riker 
and Baldwin, Jour. Bact., 28, 1934, 571.) 

Growth occurs in special alkaline media 
of pH 11.0 to 12.0 (Hofer, Jour. Amer. 
Soc. Agron.,^7, 1935,228). 

Hydrogen sulfide produced if grown in 
ZoBell and Feltham's medium (Jour. 
Bact., 28, 1934, 169). 

Distinctive characters : Browning of 
mannitol-calcium-glycerophosphate agar. 
Inability to cause plant disease or to 
produce nodules on roots of legumes. 
Complete utilization (disappearance of 
nitrate) in the peptone-salt medium of 
Riker et al. (Jour. Agr. Res., 41, 1930, 
529) and failure to absorb congo red 
(ibid., 528). 

The species bears at least superficial 
resemblances to certain Rhizobium spp., 
but may be distinguished from them by 
the first two characters listed above, and 
the following in addition : Growth at a 
reaction of pH 11-12. Heavy ring or 
pellicle formation on veal infusion broth. 
H2S production in the mannitol-tryptone 
medium of ZoBell and Feltham (loc. cit.). 
Production of milky white precipitate 
on nitrate-glycerol-soil-extract agar. 

Source: Isolated from soil. 

Habitat: Soil, around the roots of 
plants, especially legumes. 

Note: Palacios and Bari (Proc. Indian 
Acad. Sci., 3, 1936, 362; Abs. in Cent. f. 
Bakt., II Abt., 95, 1937, 423) have de- 
scribed Bacillus concomitans as a sym- 
biont from legume nodules that has no 
power to fix nitrogen although it is very 
much like legume nodule bacteria {Rhizo- 
bium spp.). This organism resembles 
Agrobaclerium radiobacter . 



Appendix: The following species prob- 
ably belong in Agrobaclerium, but are not 
sufficiently well described to make their 
relationship certain. 

1 . Agrobacterium gypsophilae (Brown) 
Starr and Weiss. {Bacterium gypsophilae 
Brown, Jour. Agr. Res., 48, 1934, 1109; 
Pseudomonas gypsophilae Stapp, Bot. 
Rev., 1, 1935, 407; Phytomonas gypso- 
philae Stapp, ibid., 407; Starr and Weiss, 
Phytopath., 33, 1943, 316.) From M. L., 
Gypsophila, a generic name. 

Rods: 0.2 to 0.8 by 0.4 to 1.4 microns. 
Motile with 1 to 4 flagella. Capsules. 
Gram-negative. 

Gelatin : Liquefaction slow, beginning 
after 1 month. 

Beef -infusion agar colonies : Circular, 
Naples yellow, smooth or rough, butyrous. 

Broth : Turbid in 24 hours. 

Milk: Coagulation and peptonization. 

Nitrites are produced from nitrates. 

Indole not produced. 

Hydrogen sulfide : A trace may be 
produced. 

Acid but not gas from glucose, sucrose, 
maltose, mannitol and glycerol. No acid 
from lactose. 

Starch not hydrolyzed. 

Aerobic, facultative. 

Distinctive characters: Differs from 
Xanthomonas beticola in starch hydrol- 
ysis, HoS production, and will not cross- 
inoculate with this species. 

Source : Isolated from several galls on 
Gypsophila. 

Habitat : Produces galls in Gypsophila 
panicvlata and related plants. 

2. Bacterium pseudotsugae Hansen and 
Smith. (Hansen and R .E. Smith, Hil- 
gardia, 10, 1937, 576; Phytoinonas pseu- 
dotsugae Burkholder, in Manual, 5th ed., 
1939, 209.) From M. L., Pseudotsuga, 
a generic name. 

Rods : 0.5 to 1.5 by 1.9 to 3.9 microns. 
Probably motile ; type of flagellation 
doubtful. Gram-negative. 

Gelatin : Liquefied. 



FAMILY RHIZOBIACEAE 



231 



Nutrient agar slant: Growth scanty, 
flat, glistening, smooth, translucent, 
whitish. 

Broth: Growth slight. Xo sediment. 

Milk: No acid. 

Nitrites produced from nitrates. 

Hydrogen sulfide production slight. 

Acid but not gas from glucose, fruc- 



tose, galactose and maltose. No acid or 
gas from lactose, sucrose or glycerol. 

Starch not hydrolyzed. 

Facultative aerobe. 

Source : Isolated from galls on Douglas 
fir in California. 

Habitat : Pathogenic on Douglas fir, 
Pseudoisuga taxifolia. 



Genus III. ChiOTD.oha.cteTiuia Bergonzini.* 

(Ann. Societa d. Naturalisti in Modena, Ser. 2, 14, 1881, 153.) Greek, chroma, 
color; M. L., bacterium, a small rod. 

Rods, 0.4 to 0.8 by 1.0 to 5.0 microns. Motile with 1 to 4 or more flagella. Gram- 
negative. A violet pigment is formed which is soluble in alcohol, but not in water or 
chloroform. Grow on ordinary culture media, usually forming acid from glucose, 
sometimes from maltose, not from lactose. Gelatin is liquefied. Indole is not pro- 
duced. Nitrate usually reduced to nitrite. Optimum temperature 20-25°C. but 
some grow well at 37°C. Usually saprophytic soil and water bacteria. 

The type species is Chromobacterium violaceum (Schroeter) Bergonzini. 

Key to the species of genus Chromobacterium. 

I. Motile rods. Single flagellum. 

A. Acid from glucose and maltose. No acid from sucrose. Nitrites produced 
from nitrates. No growth at 37°C. 

1. Chromobacterium violaceum. 
II. Motile rods. Flagella generally peritrichous. 

A. Acid from glucose. Nitrites generallj' not produced from nitrates. Good 

growth at 37°C. 

2. Chromobacterium ianthinum. 

B. Generally no acid from glucose. Nitrites produced from nitrates. No 

growth at 37°C. 

3. Chromobacterium amethystinum. 



1. Chromobacterium violaceum 

(Schroeter) Bergonzini. {Bacteridium 
violaceum Schroeter, Beitrage z. Biol. d. 
Pflanzen, 1, Heft 2, 1872, 126; Micro- 
coccus violaceus Colin, Beitrage z. Biol, 
d. Pflanzen, /, Heft 2, 1872, 157; Cromo- 
bacterium violaceum (sic) Bergonzini, 
Ann. Societa d. Naturalisti in Modena, 
Ser. 2, 14, 1881, 153; Bacillus violaceus 
Schroeter, Kryptogamen-Flora von Schle- 
sien, 3, 1886, 157; Streptococctis violaceus 
Trevisan, I generi e le specie delle 



Batteriacee, 1889, 31 ; Pseudomonas 
violacea Migula, Arb. a. d. Bakt. Inst. 
Karlsruhe, 1, 1894, 237; Bacterium vio- 
laceum Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., 2, 1896, 58; see 2 Aufl., 2, 
1899, 262.) From Latin, violaceus, 
violet-colored. 

Note: Bacterium ianthinum Zopf (Die 
Spaltpilze, 1885, 68) has been regarded as 
identical with the above organism 
by Schroeter (Kryptogamen-Flora von 
Schlesien, 3, 1, 1886, 157), and by Leh- 



* Adapted by Prof. Robert S. Breed, New York State Experiment Station, Geneva, 
New York from Cruess-Callaghan and Gorman, Scientific Proc. Royal Dublin Society, 
21, 1935, 213 in Jan. 1938; further revision, July, 1945 by Robert S. Breed with the 
assistance of Capt. W. C. Tobie, Sn. C, Old Greenwich, Conn. 



232 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



mann and Neumann (Bakt. Diag., 1 
Aufl., 2, 1896, 266; also 7 Aufl., 2, 1927, 
463). Lehmann and Neumann {loc. cil.) 
also consider Bacillus violaceus laurenti- 
cus Lustig (Diagnostik der Bakterien des 
Wassers, 1893, 103) as being identical 
with Bacterium violaceum. 

Slender rods: 0.8 to 1.0 by 2.0 to 5.0 
microns, occurring singly and in chains. 
Motile, with a single flagellum. Gram- 
negative. 

Gelatin colonies : Circular, gray, entire 
margin, assuming a violet color in the 
center. 

Gelatin stab : Infundibuliform lique- 
faction with violet sediment in fluid. 

Agar colonies : Whitish, flat, glistening, 
moist, becoming violet. 

Agar slant: Deep, violet, moist, shiny 
spreading growth. 

Broth: Slightly turbid, with violet 
ring and ropy sediment. 

Litmus milk: Violet pellicle. Diges- 
tion. Alkaline. 

Potato : Limited, dark violet growth. 

Loffler's blood serum : Slowly liquefied. 

Indole not formed. 

Nitrites produced from nitrates. 

Acid from glucose and usually from 
maltose. No acid from lactose or sucrose. 

Aerobic, facultative. 

Optimum, temperature 25° to 30°C. 
No growth at 37 °C. Slight growth at 
2° to 4°C. 

Source : Originally grown on slices of 
cooked potato exposed to air contamina- 
tion, and incubated at room temperature. 

Habitat : Water. 

2. Chromobacterium ianthinum (Zopf) 
Holland. {Bacterium ianthinum Zopf, 
Die Spaltpilze, 2 Aufl., 1884, 62; Bacillus 
janthinus Fliigge, Die Mikroorganismen, 
1886, 291 ; Bacieridium ianthinum Schroc- 
ter, Kryptogamen Flora von Schlesien, 
3, 1, 1886, 157; Pseudomonas ianthina 
Migula, Syst. d. Bakt., 2, 1900, 941 ; Pseu- 
domonas janthina Chester, Man. Determ. 
Bact., 1901, 317; Holland, Jour. Bact., 5, 
1920, 222.) From Greek, ianthinus, 
violet-blue. 



Rods : 0.5 to 0.8 by 1.5 to 5.0 microns, 
occurring singly. Motile with peritri- 
chous flagella. Gram-negative. 

Gelatin colonies: Circular, yellow, 
becoming violet. 

Gelatin stab : White to violet surface 
growth. Infundibuliform liquefaction. 

Agar colonies : Creamy center, violet 
margin. 

Agar slant : Yellowish, moist, glisten- 
ing, becoming deep violet. 

Broth: Turbid, with light violet pel- 
licle. 

Litmus milk: Slow coagulation with 
violet cream layer. Litmus decolorized 
from below. 

Potato : Violet to violet-black, spread- 
ing growth. 

Indole not formed. 

Nitrites generally not produced from 
nitrates. 

Acid from glucose. No acid from 
maltose, lactose and sucrose. 

Aerobic, facultative. 

Optimum temperature 30°C. Grows 
well at 37°C. No growth at 2 to 4°C. 

Source : Originally grown on pieces of 
pig's bladder floated on badly contam 
inated water. 

Habitat : Water and soil. This may 
be the species that causes a fatal septi- 
cemia in animals and man. See Chromo- 
bacterium violaceum manilae. 

3. Chromobacterium amethystinum 

(Chester) Holland. {Bacillus membra- 
naceus amethystinus Eisenberg, Bakt. 
Diag., 1891, 421; Bacterium amethystinus 
Chester, Ann. Rept. Del. Col. Agr. Exp. 
Sta., 9, 1897, 117; Bacterium membrana- 
ceus amethystinus Chester, ibid., 13S: 
Bacillus amethystinus Holland, Jour. 
Bact., 5, 1920, 217; not Bacillus amethys- 
tinus Chester, loc. cit., 262; Holland, 
loc. cit., 222; Bacterium memhranaceum 
amethystinum Lehmann and Neumann, 
Bakt. Diag., 7 Aufl., 2, 1927, 463; Bac- 
teriujn violaceum amethystinum Cruess- 
Callaghan and Gorman, Sci. Proc. Royal 
Dublin Society, 21, 1935, 219.) From 
Greek, bluish-violet, amethyst. 



FAMILY RHIZOBIACEAE 



233 



Rods: 0.5 to 0.8 by 1.0 to l.-l microns, 
occurring singlj'. ^lotile with a single 
or occasionally with peritrichous flagella. 
Gram-negative. 

Gelatin colonies: Thin, bluish, becom- 
ing violet, crumpled. 

Gelatin stab: Heavy, violet-black pel- 
licle. Liquefied. 

Agar colonies: Deep violet, surface 
rugose. 

Agar slant: Thick, moist, yellowish- 
white, becoming violet with metallic 
luster. 

Broth: Pellicle with violet sediment, 
fluid becoming violet. 

Litmus milk: Violet pellicle. Diges- 
tion turning alkaline. 

Potato: Deep violet, rugose spreading 
growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Usually no acid from glucose, maltose 
and sucrose. No acid from lactose. 

Aerobic, facultative. 

Optimum temperature 30°C. No 
growth at 37°C. Good growth in 7 days 
at 2 to 4°C. 

Original source : Found once by Jolles 
in spring water from Spalato. 

Habitat: Water. 

Appendix: The following organisms 
have been assigned to this genus or are 
believed to belong here. Additional 
comparative studies are badly needed. 

Bacillus cyaneo-fuscus Beijerinck. 
(Beijerinck, Bot. Ztung., .',9, 1891, 704; 
Bacterium ci/anojuscus Chester, Ann. 
Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 
116 and 132.) From black glue, blue 
Edam cheese, water and soil. 

Bacillus lacyyius Schroeter. (Schroeter 
in Cohn, Kryptogamen-Flora von Schle- 
sien, 3, 1, 1889, 158.) In greenhouse on 
fresh paint. 

Bacillus lilacinus Mace. (Traite Pra- 
tique Bact., 6« ed., 2, 1913, 416.) From 
water. 

Bacillus tuembranuceus umethystinus 
inobilis Germano. (Gcrmano, Cent. f. 
Bakt., 12, 1892, 516; Bacillus amethysti- 



nus mobilis Kruse, in Fliiggc, Die 
Mikroorganismen, 3 Aufl., 2, 1896, 313; 
Bacterium amethystinus mobilis Chester, 
Ann. Kept. Del. Col. Agr. Exp. Sta.^ 
9, 1897. 117; Bacteria?/! memhranaceus 
mobilis Chester, ibid.. 138.) Pseudo- 
monas amethyslina Migula, Syst. d. 
Bakt., 2, 1900, 944; Bacillus amethys- 
tinus Chester, Man. Determ. Bact.. 1901, 
262.) From dust. 

Bacillus pavoninus Forster. (Forster, 
in van der Sleen, Sur I'examen bacterio- 
logique qualitatif de I'eau. Arch. Teyler, 
Ser. 2, Tome 4, 3 partie, 1894, No. 59, 
Haarlem, Heritiere Loosjes. Also see 
Godfrin, These, Nancy, 1934, 46.) 
Causes blue discoloration of Edam cheese. 

Bacillus polychromogoies Chamot and 
Thiry. (Bacille polychrome, Thiry, 
Compt. rend. Soe. Biol., Paris, 48, 1896, 
885; Chamot and Thiry, Bot. Gaz., 30, 
1900, 378.) From well water at Nancy. 
Probably a Pseudomonas (Tobie, per- 
sonal communication). 

Bacillus violaceus Frankland and 
Frankland. (Frankland and Frankland, 
Ztschr. f . Hyg., e, 1888, 394 ; Pseudomonas 
pseudoianthina IMiguIa, Syst. d. Bakt., 
2, 1900, 942.) Isolated from tap water. 
Said to produce spores. 

Bacillus violaceus laurentius Jordan. 
(Jordan, Mass. State Bd. Health Rept., 
1890. 838; Bacterium violaceus laurentius 
Chester, Ann. Rept. Del. Col. Agr. 
Exp. Sta., .9, 1897, 117; Pseudomonas 
laurentia Migula, Syst. d. Bakt., 2, 
1900, 944; Bacillus violaceus Chester, 
Man. Determ. Bact., 1901, 262; Chromo- 
bacterium violaceum laurentium Ford, 
Textb. Bact., 1927, 470.) Isolated from 
sewage effluent. 

Bacillus violaceus lutetiensis Kruse. 
(Kruse, in Fliigge, Die Mikroorganismen, 
3 Aufl., 2, 1896, 311; Bacillus lutetiensis 
Chester, Man. Determ. Bact., 1901, 306; 
Chromobacterium violaceum lutetiense 
Ford, Textb. Bact., 1927, 470.) From 
water. 

Bacillus violaceus sarloryi Waeldele. 
(These, Pharm. Strasbourg, 1938, 55.) 



234 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



From dental pus. Said to form spores. 

Bacterium cristallino violaceum Chol- 
kevitch. (Cholkevitch, 1922, quoted 
from Godfrin, Contribution a I'etude des 
bacteries bleues et violettes. Th6se, 
Nancy, 1934, 93.) From peat. 

ChromobacteriimibamptoniiJieTgey et a\ . 
{Bacillus memhranacens arnethystinus II, 
Bampton, Cent. f. Bakt., I Abt., Orig., 
71, 1913, 137; Bergey et al.. Manual, 1st 
ed., 1923, 119; Chromobacteriutn mem- 
branaceum amethystinum II Ford, Textb. 
Bact., 1927, 473.) From water. 

Chromobacterium coeruleum (Voges) 
Bergey et al. (Bacillus coeruleus Voges, 
Cent. f. Bakt., U, 1893, 303; Bacterium 
coeruleus Chester, Ann. Rept. Del. Col. 
Agr. Exp. Sta.,9, 1897, 117 ; Pseudomonas 
coerulea Migula, Syst. d. Bakt., 2, 1900, 
945; Bergey et al.. Manual, 1st ed., 1923, 
120.) From water. 

Chromobacterium cohaerens Grimes. 
(Sci. Proc. Royal Dublin Society, 19, 
1930,381.) From well water. 

Chromobacterium hibernicum Grimes. 
(Sci. Proc. Royal Dublin Society, 19, 
1930,381.) From well water. 

Chromobacterium lividum (Voges) Hol- 
land. (Plagge and Proskauer, Zeitsch. 
f. Hyg., 2, 1887, 463; Bacillus lividus 
Voges, Cent. f. Bakt., U, 1893, 303; 
relationship to Bacillus lividus Zimmer- 
mann uncertain. Die Bakt. unserer 
Trink- und Nutzwasser, Chemnitz, 2, 
1894, 18; Bacillus violaceus berolineiisis 
Kruse, in Flugge, Die Mikroorganismen, 
3 Aufl., 2, 1896, 311; Bacterium lividus 
Chester, Ann. Rept. Del. Col. Agr. Exp. 
Sta., 9, 1897, 117; Bacillus berolinensis 
Chester, Man. Determ. Bact., 1901, 305; 
Holland, Jour. Bact., 5, 1920, 215.) 
From water. 

Chromobacleriuin maris-mortui Ela- 
zari-Volcani. (Studies on the Micro- 
flora of the Dead Sea, Thesis, Hebrew 
Univ., Jerusalem, 1940, vii and 76.) 
From the Dead Sea. 

Chromobacterium membranacemn Ber- 
gey et al. (Bacillus membranaceus ame- 
thystinus I, Bampton, Cent. f. Bakt., I 
Abt., Orig., 71, 1913, 135; Bergey et al., 
Manual, 1st ed., 1923, 119; Chromobac- 



terium membranaceum amethystinum I 
Ford, Textb. Bact., 1927, 472.) From 
water. 

Chromobacterium membranaceum ame- 
thystinum III Ford. (Ford, Textb. 
Bact., 1927, 474; Bacillus membranaceus 
arnethystinus III Bampton, Cent. f. 
Bakt., I Abt., Orig., 7i, 1913, 138.) From 
water. 

Chromobacterium membranaceum ame- 
thystinum IV Ford. (Ford, Textb. 
Bact., 1927, 474; Bacillus membranaceus 
arnethystinus IV Bampton, Cent. f. 
Bakt., I Abt., Orig., 71, 1913, 138.) From 
water. 

Chromobacterium smithii (Chester) 
Bergey et al. (Bacillus coeruleus Smith, 
Medical News, 2, 1887, 758; Bacterium 
coeruleus Chester, Ann. Rept. Del. Col. 
Agr. Exp. Sta., 9, 1897, 118; Pseudomonas 
smithii Chester, Man. Determ. Bact., 
1901, 318; Chromobacterium coeruleum 
Ford, Textb. Bact., 1927, 475; not Chro- 
mobacterium coeruleum Bergey et al., 
Manual, 1st ed., 1923, 120; Bergey et al., 
ibid., 121.) From water. 

Chromobacterium violaceum manilae 
Ford. (Bacillus violaceus manilae Wool- 
ley, U. S. Dept. Int., Bur. Govt. Labs. 
Bull. 15, 1904 and Bull. Johns Hopkins 
Hosp., 16, 1905, 89; Ford, Textb. Bact., 
1927, 471.) Isolated from fatal septice- 
mias in water buffalo (Woolley) and man 
(Schattenberg and Harris, Jour. Bact., 
U, 1942,509). More likely to be a variety 
of Chromobacterium ianthinum which 
grows at 37°C. than of C. violaceum which 
does not grow at 37°C. 

Chromobacterium viscofucatum (Harri- 
son and Barlow) Bergey et al. (Bacterium 
viscofucatum and Bacillus viscofucatus 
Harrison and Barlow, Cent. f. Bakt., II 
Abt., 15, 1905, 517; Trans. Royal Soc. 
Canada, 2nd Ser., 11, 1905; Bergey et al.. 
Manual, 1st ed., 1923, 119.) From oily 
butter. Probably a non-motile Pseudo- 
monas (Tobie, personal communication). 

Chromobacterium viscosum Grimes. 
(Cent. f. Bakt., II Abt., 72, 1927, 367.) 
From butter. 

Pseudomonas pseudoviolacea Migula. 
(Syst. d. Bakt., 2, 1900, 943.) From 
river water. 



FAMILY MICROCOCCACEAE 235 

FAMILY V. MICROCOCCACEAE PRIBRAM.* 
(Jour. Bact., 18, 1929, 385.) 

Cells without endospores except in Sporosarcina. Cells in their free condition 
spherical; during division somewhat elliptical. Division in two or three planes. If 
the cells remain in contact after division, they are frequently flattened in the plane 
of last division. They occur singly, in pairs, tetrads, packets or irregular masses. 
Motility rare. Generally Gram-positive. Many species form a yellow, orange, pink 
or red pigment. Most species are preferablj^ aerobic, producing abundant growth on 
ordinar}^ culture media, but capable of slight anaerobic growth. A few species are 
strictly anaerobic. ^Metabolism heterotrophic. Carbohydrates are frequently fer- 
mented to acid. Gelatin is often liquefied. Facultative parasites and saprophytes. 
Frequently live on the skin, in skin glands or skin gland secretions of Vertcbrata. 

Key to the genera of family Micrococcaceae. 

I. Cells occur in plates, groups or in irregular packets and masses, never in chains. 
Pigment, when present, is yellow, orange or red. Gram-positive to Gram- 
negative. 

Genus I. Micrococcus, p. 235. 
II. On the animal body and in special media cells occur as tetrads. In ordinary 
media cells may occur in pairs and irregular masses. White to pale yellow. 

Genus II. Gaffkya, p. 283. 
III. Cells occur in regular packets. Yellow or orange pigment usually formed. 

Genus III. Sarcina, p. 285. 

Genus I. Micrococcus Cohn.* 

(Cohn, Beitrage z. Biol. d. Pflanzen, /, Heft 2, 1872, 153; Microsphaera Cohn, 
Arch. f. path. Anat., 55, 1872, 237; not Microsphaera Leveille, Ann. Sci. Nat. Bot., 
Ser. 3, 16, 1851, 381 ; Ascococcus Cohn, Beitrage z. Biol. d. Pflanzen, 1, Heft 3, 1875, 
154; Pediococcus Balcke, Wchnschr. f. Brauerei, 1, 1884, 183; Merista Van Tieghem, 
Traite de Botanique, Paris, 1884, 1114; Staphylococcus Rosenbach, Mikroorganismen 
bei den Wundinfektions-krankheiten des Menschen, 1884, 27; Monococcns Miller, 
Deutsch. med. Wchnschr., 12, 1886, No. 8, 117; Botryomyccs Bollinger, Deutsch. 
Ztschr. f. Tiermed., 13, 1887, 77; Urococcus Miquel, Ann. Microg., /, 1888, 518; Galac- 
tococcus Guillebeau, Jahrb. d. Schweiz, 4, 1890, 32; Rhudococcus Zopf, Ber. d. deutsch. 
Bot. Gesellsch., Berlin, 9, 1891, 28; Pyococcus Ludwig, Lehrb. d. niederen Kryptog., 
1892, 27; Planococcus Migula, Arb. Bakt. Inst. Karlsruhe, 1, 1894, 236; Carphococcus 
Hohl, Cent. f. Bakt., II Abt., 9, 1902, 338; .4Z6ococcws Winslow and Rogers, Jour. Inf. 
Dis., 3, 1906, 541; Aurococcus Winslow and Rogers, ibid., 540; Pedioplana Wolff,. 
Cent. f. Bakt., II Abt., 18, 1907,9; Melococcus Nedrigailov, Charkov Med. Zurnal, 4, 
1907, 301; Solidococcus, Liquidococcus, Indolococcus and Peptonococcus Orla-Jensen, 
Cent. f. Bakt., II Abt., 22, 1909, 332; Planomerista Vuillemin, Ann. Mycol., 
11, 1913, 525; Tetracoccus Orla-Jensen (in part), The Lactic Acid Bacteria, 1919, 76.) 
From Greek micrus, small; coccus, a grain; M. L., a sphere. 

Cells in plates or irregular masses (never in long chains or packets). Gram-posi- 
tive to Gram-negative. Growth on agar usually abundant, some species form no 

* The genera Micrococcus and Staphylococcus have been combined and completely 
revised by Prof. G. J. Hucker, New York State Experiment Station, Geneva, New 
York, March, 1943 so far as the aerobic species are concerned. Dr. Ivan C. Hall, 
Presbyterian Hospital, New York City, revised the anaerobic section, January, 1944. 



236 MANUAL OF DETERMINATIVE BACTERIOLOGY 

pigment but others form yellow or less commonly orange, or red pigment. Glucose 
broth slightly acid, lactose broth generally neutral. Gelatin frequently liquefied, 
but not rapidly. Facultative parasites and saprophytes. 
The type species is Micrococcus luieus (Schroeter) Cohn. 

Key to the species of genus Micrococcus. 
1. Aerobic to facultative anaerobic species. 
I. No pink or red pigment on agar media. 

A. Nitrites not produced from nitrates. 

1. Utilize NH4H2PO4 as sole source of nitrogen.* 

--. - a. Yellow pigment on agar media. Not acido-proteolytic, 

1. Micrococcus liiteus. 
aa. No pigment produced. Not acido-proteolytic. 

b. Utilizes urea as a sole source of nitrogen.** 

2. Micrococcus ureae. 
bb. Does not utilize urea. 

3. Micrococcus freudenreichii. 
aaa. Acido-proteolytic in litmus milk. 

8. Micrococcus caseolyticus. 

2. Do not utilize NH4H2PO4 as sole source of nitrogen. 

a. Yellow pigment produced. 

4. Micrococcus flavus. 
aa. No pigment produced. 

5. Micrococcus candidus. 

B. Nitrites produced from nitrates. 

1. Utilize NH4H2PO4 as sole source of nitrogen. 

a. Yellow pigment on agar media. Not acido-proteolytic. 
b. Gelatin liquefied. 

6. Micrococcus conglomeratus . 
bb. Gelatin not liquefied. 

7. Micrococcus varians. 

aa. Usually not chromogenic. Actively acido-proteolytic inlitmus milk. 

S. Micrococcus caseolyticus. 

2. Do not utilize NH4H2PO4 as sole source of nitrogen. 

a. Gelatin liquefied. Ferment mannitol. 
b. Abundant orange growth on agar media. 

9a. Micrococcus pyogenes var. aureus. 
bb. Abundant white growth on agar media. 

9b. Micrococcus pyogenes var. albus. 
bbb. Yellow growth on agar media. 

10. Micrococcus citreus. 
aa. Gelatin not liquefied or very slowly liquefied. 

b. Abundant orange to white growth on agar media. Ferments 
mannitol. 

11. Micrococcus aurantiacus. 

bb. Scant white translucent growth on agar media. Does not 
ferment mannitol. 

12. Micrococcus epidermidis. 

* That is, will grow and produce acid (sometimes slowly) on slants containing 
1.5 per cent washed agar, 0.1 per cent ammonium phosphate, 1.0 per cent glucose, 
0.02 per cent potassium chloride, 0.02 per cent magnesium sulfate. Add brom-cresol- 
purple as an indicator (Hucker, N. Y. State E.xper. Sta., Tech. Bui. 100, 1924, 25; 
Tech. Bui. 101, 1924, 36-40) ; Manual Pure Culture Study of Bacteria. Soc. Amer. 
Bact., Geneva, N. Y., Leaflet II, 9th ed., 1944, 14.) 



FAMILY MICKOCOCCACEAE 



237 



II. Pink or red pigment on agar media. 

A. Gelatin liquefied, slowly. Produces rose-colored pigment. 

13. Micrococcus roseus. 

B. Gelatin not liquefied. 

1. Non-motile. 

a. Produces cinnabar-colored pigment on gelatin. 

14. Micrococcus cinnabareus. 

aa. Produces light, flesh-colored pigment on agar slant. Ferments 
glycerol and mannitol. 

15. Micrococcus rubens. 

aaa. Produces brick-colored pigment on agar slant. Does not ferment 
glycerol and mannitol. 

16. Micrococcus rhodochrous. 

2. Motile. Produces red pigment. 

17. Micrococcus agilis. 
2. Anaerobic species. 

I. Forms gas from nitrogenous media. 

A. Acid from glucose. 

18. Micrococcus aerogenes. 

B. No acid from glucose. 

1. No blackening of colonies in deep agar. 

19. Micrococcus asaccharolyiicus. 

2. Hj'drogen sulfide formed. Deep agar colonies become black. 

20. Micrococcus niger. 
II. No gas formed from nitrogenous media. 

A. Acid from glucose. 

1. Acid from lactose. 

21. Micrococcus grigoroffi. 

2. No acid from lactose. 

22. Micrococcus anaerobius. 



1. Micrococcus luteus (Schroeter) 
Cohu. {Bactcridium lutcum Schroeter, 
Beitr. z. Biol. d. Pflan., /, Heft 2, 1872, 
119; Cohn, ibid., 153.) From Latin, 
luicus golden-yellow. 

Spheres: 1.0 to 1.2 microns, occurring 
in pairs and fours. Non-motile. Gram- 
positive. 

Gelatin colonies: Yellowish-white to 
yellow, raised, with undulate margin. 

Gelatin stab: No liquefaction. 

Agar colonies : Small, yellowish, glisten- 
ing, raised. 

Agar slant: Citron-yellow, smooth. 

Broth : Clear, with yellowish sediment. 

Litmus milk: Usually slighth- acid, 
not coagulated. 

Potato: Thin, glistening, citron-yellow 
growth. 

Indole not formed. 

Nitrites not produced from nitrates. 



Acid from glucose, sucrose and man- 
nitol. No acid from lactose. 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

Utilizes NH4H2 PO4 as a source of 
nitrogen. 

Saprophytic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Isolated by Schroeter from 
dust contaminations on cooked potato. 

Habitat : Found in skim milk and dair}^ 
products, and on dust particles. 

2. Micrococcus ureae Cohn. (Cohn, 
Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 
1872, 158; not Micrococcus ureae Fliigge, 
Die Mikroorganismen, 2 Aufl., 1886, 
169; Merisla ureae Prazmowski, Biol. 
Cent., 8, 1888, 301; Streptococcus ureae 
Trevisan, I generi e le specie delle Bat- 



238 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



teriacee, Milan, 1889, 31 ; Urococcus 
ureae Beijerinck, Cent. f. Bakt., II Abt., 
7, 1901, 52; Albococcus ureae Kligler, Jour. 
Infect. Dis., 13, 1943, 442; Staphylococcus 
wreae Holland, Jour. Bact., 5, 1920, 225.) 
From Greek, urum, urine; M. L., urea, 
urea. 

See Micrococcus liquefaciens Migula 
in the appendix for references to the 
gelatin-liquefying form of the species. 

Spheres: 0.8 to 1.0 micron, occurring 
singly, in pairs and in clumps. Never in 
chains. Non-motile. Gram-variable. 

Gelatin colonies : Small, white, translu- 
cent, slimy, becoming fissured. 

Gelatin stab: Slight, white growth. 
Very slow or no liquefaction. 

Agar colonies: White, slightly raised. 

Agar slant: Grayish -white, raised, 
glistening, butyrous. 

Broth: Turbid, with viscid sediment. 

Litmus milk : Slightly alkaline ; litmus 
slowly reduced. 

Milk: Acid. 

Potato: Slight, grayish to pale olive 
growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Urea fermented to ammonium car- 
bonate. 

Acid produced from glucose, lactose, 
sucrose and mannitol. 

Starch not hydrolyzed. 

Ammonium salts are utilized. 

Ammonia produced from peptone. 

Saprophytic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Isolated from fermenting urine. 

Habitat : Found in stale urine and in 
soil containing urine. 



Sternberg, Man. of Bact., 1893, 601; 
Micrococcus acidilactis Migula, Syst. d. 
Bakt., 2, 1900, 112; Micrococcus acidifi- 
cans Migula, ibid.); Micrococcus lactis 
viscosus Sternberg, Man. of Bact., 1893, 
604; Micrococcus amarijaciens Migula, 
Syst. d. Bakt., 2, 1900, 100; Coccus lactis 
viscosi Gruber, Cent. f. Bakt., II Abt., 
9, 1902, 790 {Micrococcus lactis viscosi 
Lohnis, Cent. f. Bakt., II Abt., 18, 1907, 
144) ; Micrococcus lactis albidus Conn, 
Esten and Stocking, Storrs Agr. Exp. 
Sta. 18th Ann. Rept., 1906, 91. 

Spheres: 2.0 microns in diameter, oc- 
curing singly and in clumps, rarely in 
short chains. Non-motile. Gram-posi- 
tive. 

Milk gelatin colonies: Small, white, 
opaque. 

Milk gelatin stab : Inf undibuliform 
liquefaction. 

Agar colonies: White, slimy. 

Agar streak: White, smooth. 

Broth : Turbid, with white sediment. 

Litmus milk: Acid; coagulated; pep- 
tonized. 

Potato: Moderate white to yellow 
streak. 

Indole not formed. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

Does not utilize urea as a source of ni- 
trogen. 

Acid from glucose, lactose and sucrose. 
Some strains form acid from mannitol ; 
others from glycerol. 

Saprophytic. 

Aerobic. 

Optimum temperature 20°C. 

Habitat : Milk and dairy utensils. 



3. Micrococcus freudenreichii Guille- 
beau. (Landwirtsch. Jahrb. d. Schweiz, 
5, 1891, 135.) Named for E. v. Freuden- 
reich, Swiss bacteriologist. 

Synonyms: Micrococcus acidi lactis 
Krueger, Cent. f. Bakt., 7, 1890, 464 
{Micrococcus acidi lactis liquefaciens 
Eisenberg, Bakt. Diag., 3 Aufl., 1891, 
409 ; Micrococcus acidi lactici liquefaciens 



4. Micrococcus flavus Trevisan. {Mi- 
crococcus flavus liquefaciens Fliigge, Die 
Mikroorganismen, 2 Aufl., 1886, 174; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 34; Micrococcus 
flavus-liquefaciens Chester, Man. De- 
term. Bact., 1901, 99.) From Latin, 
flavus, yellow. 

Spheres: 0.8 to 0.9 micron, occurring 



FAMILY MICROCOCCACEAE 



239 



singly, in clumps, and occasionally in 
fours. Occasionally cultures are found 
that are motile with a single flagellum. 
Otherwise non-motile. Gram-variable. 

Gelatin colonies: Small, circular, yel- 
lowish to yellowish-brown, somewhat 
serrate margin, granulated, sharply con- 
toured. 

Gelatin stab: Yellow, wrinkled surface 
growth with slow, crateriform lique- 
faction. 

Agar colonies: Small, pale yellowish, 
homogeneous, entire. 

Agar slant : Canary -yellow, somewhat 
dry, wrinkled, raised, entire. 

Broth : Turbid with yellowish ring and 
sediment. 

Litmus milk: Slightly acid, soft 
coagulum formed, with slight reduction; 
slowly peptonized. 

Potato: Slight, canary-yellow growth. 

Indole is not formed. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Acid is generally formed from glucose 
and lactose. Sucrose, glycerol and man- 
nitol generally not fermented. 

Ammonium salts are utilized. 

Ammonia produced from peptone. 

Non-pathogenic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Original source not given. 

Habitat : Found in skin gland secre- 
tions, milk, dairy products, and dairy 
utensils. 

5. Micrococcus candidus Cohn. 
(Cohn, Beitr. z. Biol. d. Pflanzen, 1, 
Heft 2, 1872, 160 ; Staphylococcus candidus 
Holland, Jour. Bact., 5, 1920, 223.) From 
Latin candidus, shining white. 

Spheres: 0.5 to 0.7 micron, occurring 
singly. Non-motile. Gram-positive. 

Gelatin colonies : White, granular, with 
irregular or entire margin. 

Gelatin stab: White surface growth. 
Filiform. No liquefaction. 

Agar colonies: Punctiform, white, 
smooth, entire, iridescent. 



Agar slant: Smooth, white, glistening, 
iridescent. 

Broth: Turbid, with pellicle. 

Litmus milk : Slightly acid ; not coagu- 
lated. 

Potato: Thick, porcelain white, 
glistening. 

Indole not produced. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

Ammonium salts not utilized. 

Acid from glucose, sucrose, lactose 
and glycerol. 

Non-pathogenic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Originally appeared as white 
colonies on cooked potato exposed to dust 
contaminations. 

Habitat: Found in skin secretions, 
milk and dairy products. 

6. Micrococcus conglomeratus Migula. 
(Citronengelber Diplococcus, Bumm, 
Der Mikroorganismen der gonorrhoischen 
Schleimhauterkrankungen, 1 Aufl., 1S85, 
17; Micrococcus citreus conglomeratus 
Flligge, Die Mikroorganismen, 2 Aufl., 
1886, 182; Diplococcus citreus conglomer- 
atus Bumm, ibid., 2 Aufl., 1887; Neisseria 
citrea TrevLsan, I generi e le specie delle 
Batteriacee, Milan, 1889, 32; Mcrismo- 
pcdia citreus conglomeratus Dyar, Ann, 
N. Y. Acad. Sci., 8, 1895, 352; Migula, 
Syst. d. Bakt., 2, 1900, 146; not Micro- 
coccus conglomeratus Weichselbaum, 1887, 
see Trevisan, loc. cit., 33; Micrococcus 
citreus Winslow and Winslow, Systematic 
Relationships of the Coccaceae, 1908, 
218.) From Latin, conglomeratus, rolled 
together, crowded. 

Spheres: 0.8 to 1.2 microns, occurring 
singly, in pairs, in fours, and in large 
clumps. Non-motile. Gram-variable. 

Gelatin colonies: Small, circular, yel- 
low with radiate margin. 

Gelatin stab : Slow crateriform lique- 
faction. 



240 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Agar colonies : Luxuriant, moist, sulfur 
yellow. 

Agar slant: Light yellow, plumose, 
slightly rugose, somewhat dull, raised 
center and transparent margin. 

Broth: Turbid, with light orange ring 
and sediment. 

Milk: Generally acid but not suffi- 
cient to curdle. 

Potato: No growth. 

Indole not formed. 

Nitrites produced from nitrates. 

Blood not hemolyzed. 

Starch not hydrolyzed. 

Acid from glucose and lactose gen- 
erally, sometimes from sucrose. Manni- 
tol and glycerol generally not fermented. 

Ammonia produced from peptone. 

Utilizes NH4H2PO4 as a source of 
nitrogen. 

Resistant to drying and heat. 

Non-pathogenic. 

Aerobic . 

Optimum temperature 25°C. 

Source : Found in gonorrhoeal pus and 
dust. 

Habitat: Infections, milk, dairy prod- 
ucts, dairy utensils, water, common. 

7. Micrococcus varians Migula. 
(Merismopedia Jlava varians Dyar, Ann. 
N. Y. Acad. Sci., 8, 1895, 346; Migula, 
Syst. d. Bakt., 2, 1900, Ido; Merismopedia 
flava-varians Chester, Man. Determ. 
Bact., 1901, 103; Micrococcus lactis vari- 
ans Conn, Esten and Stocking, Storrs 
Agr. Exp. Sta. Rept. for 1906, 121.) From 
Latin, varians varying. 

Spheres: 0.8 to 1.0 micron, occurring 
singly, in pairs and in fours. Occa- 
sionally cultures are found that are motile 
with a single flagellum. Otherwise non- 
motile. Gram -variable. 

Gelatin colonies: Small, circular, whit- 
ish to yellow, capitate, moruloid. 

Gelatin stab : Scant growth. No lique- 
faction. 

Agar colonies: Small, yellow, raised, 
glistening. 



Agar slant: Plumose, yellow, vari- 
egated. 

Broth: Turbid, with yellow, granular 
sediment. 

Litmus milk : Acid; coagulated on boil- 
ing. 

Potato: Raised, dry, bright-yellow, 
glistening. 

Indole not formed. 

Nitrites produced from nitrates. 

Acid from glucose, lactose, sucrose, 
raffinose and frequently from glycerol 
and mannitol. No acid from salicin or 
inulin. 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

Utilizes NH4H2PO4 as a source of 
nitrogen. 

Saprophytic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Original strains found in a 
contaminated jar of sterilized milk. 

Habitat : Has been found in body 
secretions, dairy products, dairy utensils, 
dust and water, including sea water. 

8. Micrococcus caseolyticus Evans. 
(Evans, Jour. Inf. Dis., 18, 1916, 455; 
Micrococcus casei Hucker, N. Y. Agr. 
Exp. Sta. Tech. Bull. 102, 1924, 17; Prob- 
ably Micrococcus casei Holland, Jour. 
Bact., 5, 1920, 223.) 

Identical in part with Micrococcus casei 
acidoproteolyticus I and // Gorini, Rev. 
Gen. du Lait, 8, 1910, 337; Tetracoccus 
liquejaciens Orla-Jensen, The Lactic 
Acid Bacteria, 1919, 80 {Micrococcus casei 
liquefaciens Orla-Jensen, Doktordispu- 
tats, 1904; Tetracoccus casei liquefaciens 
Orla-Jensen, The Lactic Acid Bacteria, 
1919, 80 ; Micrococcus liquefaciens Holland , 
Jour. Bact., 5, 1920, 224. Also see ref- 
erences under Streptococcus liquefaciens.) 
From Latin, caseus, cheese, casein; and 
Greek, lyticus, able to dissolve; M. L., 
dissolving, digesting. 

Spheres, variable in size, occurring in 
clumps. Non-motile. Gram-positive. 

Gelatin stab: Liquefaction generally 



FAMILY MICROOOCCACEAE 



241 



begins after first day and continues 
rapidly. 

Agar colonies : Yellow to orange (Evans, 
loc. cit.), pearly white (Hucker, loc. cit.). 

Agar stroke : Yellow to orange (Evans, 
loc. cit.), pearly white (Hucker, ?oc. cii.), 
luxuriant growth. 

Broth : Generally grows with smooth 
turbidity although certain strains give 
heavy precipitate with clear supernatant 
fluid. 

Litmus milk: Acid, peptonized. Whej- 
generally clear. 

Potato : Scanty white growth. Certain 
strains may show yellow pigment. 

Indole not formed. 

Nitrites usually produced from ni- 
trates. 

Acid from glucose, lactose, maltose, 
mannitol and glycerol. Xo action on 
raffinose. 

Forms dextrorotarj' lactic acid (Orla- 
.Tensen, 1919, loc. cit.). 

Asparagin and urea decomposed by 
some strains. 

Utilizes NH4H2PO4 as a source of nitro- 
gen. 

Optimum temperature 22°C. 

Aerobic. 

Saprophytic. 

Source : Eight cultures from bovine 
udder. 

Habitat: Milk and dairy products, 
especially cheese, dairy utensils. 

9a. Micrococcus pyogenes var. aureus 
(Rosenbach) Zopf . (Staphylococcus pyo- 
genes aureus Rosenbach, Mikroorganis- 
men bei den Wundinfectionskrankheiten 
des Menschen, Wiesbaden, 1884, 19; 
Staphylococcus aureus Rosenbach, ibid., 
27; Micrococcus pyogenes var. aureus 
Zopf, Die Spaltpilze, 3 Aufl.. 1885, 56; 
Micrococcus aureus Zopf, ibid., 57; 
Micrococcus pyogenes Lehmann and Xeu- 
mann, Bakt. Diag., 1 Aufl., 2, 1896, 165; 
Aiirococcus aureus Winslow and Rogers, 
.Jour. Inf. Dis., 3, 1906, 554; Micrococcus 
lactis varians Conn, Esten and Stocking, 
Storrs Agr. Exp. Sta. Rept. for 1906, 



121 ; Staphylococcus pyogenes Andre wes 
and Gordon, Rept. (35th) Med. Officer 
Local Govt. Board, London, 1907, 549; 
(Tetracoccus) Micrococcus pyogenes 
aureus Orla-Jensen, The Lactic Acid 
Bacteria, 1919, 81 ; Staphylococcus pyo- 
genes-aureus Holland, Jour. Bact., 5, 
1920, 225.) From Greek, pyon, pus; 
M. h., -genes, producing. From Latin, 
aureus, golden. 

Spheres: 0.8 to 1.0 micron, occurring 
singl}', in pairs, in short chains, and in 
irregular clumps. Xon-motile. Gram- 
positive. 

Gelatin stab : Saccate liquefaction with 
yellowish pellicle and yellow to orange 
sediment. 

Agar colonies: Circular, smooth, yel- 
lowish to orange, glistening, butyrous, 
entire. 

Agar slant : Abundant, opaque, smooth, 
flat, moist, yellowish to orange. 

Broth : Turbid with yellowish ring and 
sediment, becoming clear. 

Litmus milk: Acid; coagulated. 

Potato: Abundant, orange, glistening. 

Indole not formed. 

Nitrites produced from nitrates. 

Acid from glucose, lactose, sucrose, 
mannitol and glycerol, but not from 
raffinose, salicin or inulin. 

Forms inactive or levorotary lactic 
acid (Orla-Jensen, loc. cit.). 

Slight H2S formation. 

Starch not hydrolyzed. 

Does not utilize XH4H;P04, as a source 
of nitrogen. 

Ammonia produced from peptone. 

Pathogenic. Individual strains vary 
in their ability to produce hemolysin, 
coagulase and other metabolic products. 

Certain strains, under favorable con- 
ditions, produce not only exotoxins 
(hematoxin, dermatoxin, lethal toxin, 
etc.) but also a potent enterotoxin 
which is a significant cause of food poi- 
soning (Dolman and Wilson, Jour. 
Immunology, 35, 1938. 13). 

Aerobic, facultative. 

Optimum temperature 37°C. 

Source : Isolated from pus in wounds. 



242 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Habitat : Skin and mucous membranes. 
The cause of boils, abscesses, furuncles 
suppuration in wounds, etc. 

9b. Micrococcus pyogenes var. albus 
(Rosenbach) Schroeter. {Staphylococcus 
pyogenes albus Rosenbach, Mikroorganis- 
men bei den Wundinfektionskrankheiten 
des Menschcn, Wiesbaden, 1884, 2 ; Staph- 
ylococcus albus Rosenbach, ibid., 27; 
Micrococcus pyogenes var. albus Schroe- 
ter, in Cohn, Kryptog. Flora v. Schlesien, 
3, 1, 1886, 147; Micrococcus pyogenes 
Migula, Syst. d. Bakt., 2, 1900, 87; Alba- 
coccus pyogenes Winslow and Rogers, 
Jour. Inf. Dis., 3, 1906, 544; Micrococcus 
oZ6ws Buchanan, Veterinary Bacteriology 
1911, 196; (Tetracoccus) Microccocus 
pyogenes albus Orla-Jensen, The Lactic 
Acid Bacteria, 1919, 81; Staphylococcus 
pyogenes-albus Holland, Jour. Bact., 5, 
1920, 225.) From Latin, albus, white. 

Spheres: 0.6 to 0.8 micron, occurring 
singly, in pairs and in irregular groups. 
Non-motile. Gram-positive. 

Gelatin stab : Saccate liquefaction with 
heavy white sediment. 

Agar colonies : Circular, white, smooth, 
glistening, entire. 

Ten per cent evaporated milk agar: 
Growth at 20°C frequently orange 
(Chapman, Jour. Bact., 45, 1943, 405). 

Agar slant: Abundant, white, smooth, 
glistening. 

Broth: Turbid, with delicate pellicle 
and white sediment. 

Litmus milk : Acid ; coagulated. Little 
or no visible peptonization. 

Potato: Thick, smooth, white, glisten- 
ing. 

Indole not formed. 

Nitrites produced from nitrates. 

Hydrogen sulfide is formed. 

Acid formed from glucose, lactose, 
sucrose, glycerol and mannitol, but not 
from raffinose, salicin and inulin. 

Forms inactive or levorotary lactic 
acid (Orla-Jensen, loc. cit.). 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

Does not utilize NH4H2PO4 as a source 
of nitrogen. 



Pathogenic. Production of toxins, 
coagulase and hemolysin as in Micro- 
coccus aureus. 

Aerobic, facultative. 

Optimum temperature 37°C. 

Source : Originally isolated from pus. 

Habitat : Skin and mucous membranes. 
Occurs in wounds, boils, abscesses, etc. 

10. Micrococcus citreus Migula. 
(Staphylococcus pyogenes citreus Passet, 
Aetiologie der eiterigen phlegmone des 
]\Ienschen, Berlin, 1885, 9; Micrococcus 
pyogenes citreus Schroeter, in Cohn, 
Kryptog. Flora v. Schlesien, 3, 1, 1886, 
147; Migula, Syst. d. Bakt., 2, 1900, 147; 
Staphylococcus citreus Bergey et al., 
Manual, 1st ed., 1923,55.) From Latin, 
citreus, of or relating to the citrus tree; 
M. L., lemon yellow. 

Spheres: 0.9 micron, occurring singly. 
Gram-positive. 

Gelatin colonies : Circular, pale yellow, 
granular, entire, liquefying in 6 days. 

Gelatin stab : Lemon yellow surface 
growth sinking into the medium. Gray- 
ish-white growth in stab. Complete 
liquefaction in 43 days. 

Agar colonies: Small, yellow, smooth, 
entire. 

Agar slant: Broad, lemon yellow, glis- 
tening, elastic. 

Broth: Turbid, with yellow sediment 
and pellicle. 

Litmus milk: Acid, with slow coagula- 
tion. 

Potato : Thin, grayish streak, becoming 
citron yellow. 

Indole not formed. 

Nitrites produced from nitrates. 

Starch not hydrolyzed. 

Acid from glucose, lactose, sucrose, 
raffinose, inulin, salicin, glycerol and 
mannitol. 

Does not utilize NH4H2PO4 as a source 
of nitrogen. 

Ammonia produced from peptone. 

Aerobic, facultative. 

Pathogenic. 

Optimum temperature 37 °C. 

Source : Originally isolated from pus. 



FAMILY MICROCOCCACEAE 



243 



Habitat : Skin and mucous membranes 
of vertebrates. 

11. Micrococcus aurantiacus (Schroe- 
ter) Cohn. (Bacteridium aurantiacuin 
Schroeter, Beitr. z. Biol., 1, Heft 2, 1872, 
126; Colin, Beitr. z. Biol., 7, Heft 2, 1872, 
154; Pediococcus aurantiacus De Toni and 
Trevisan, in Saccardo, Sylloge Fungorum, 
8, 1889, 1051; Micrococcus aurantiacus- 
sorghi Bruyning, Arch. Neer. Sci. E.xaet. 
et Nat., 1, 1898, 297; Streptococcus 
aurantiacus Chester, Man. Determ. 
Bact., 1901, 69; Aurococcus aurantiacus 
Winslow and Winslow, Systematic Rela- 
tionships of the Coccaceae, 1908, 186; 
Sarcina aurantiaca Holland, Jour. Bact., 
5, 1920, 225 (not Sarcina aurantiaca 
Flijgge, Die Mikroorganismen, 2 Aufl., 
1886, 180); Staphylococcus aurantiacus 
Holland, ibid.) From Latin, aurum, 
gold;M. L., aurantium , the orange ;M.L., 
aurantiacus, orange-colored. 

Spheres: Slightly ellipsoidal, 1.3 to 1.5 
microns, occurring singly, in short chains 
and in small clumps. Xon-motile. 
Gram-positive. 

Gelatin colonies: Circular to oval, 
smooth, glistening with yellow to orange 
center. 

Gelatin stab : Yellow surface growth . 
No liquefaction. 

Agar colonies: Circular, smooth, glis- 
tening, yellow to orange, entire. 

Agar slant : Buff to scant orange-j^ellow, 
beaded growth, raised, glistening. 

Broth : Turbid, with pellicle. 

Litmus milk: Faintly acid, no coagu- 
lation. 

Potato : Slimy, yellow growth. Pig- 
ment is insoluble in alcohol and ether. 

Indole not produced. 

Nitrites generally produced from ni- 
trates. 

Slight acidity from glucose, fructose, 
sucrose, lactose and mannitol. Xo acid 
from raffinose, salicin, inulin. 

Starch not hydrolyzed. 

Ammonia produced from peptone. 

No growth in ammonium media. 

May be pathogenic. 

Optimum temperature 25°C. 



Aerobic. 

Source : First isolated from colonies 
that grew on boiled egg exposed to dust 
contamination. 

Habitat : Usually isolated from infec- 
tions but also found in milk, cheese and 
dust. 

Note: Albococcus cpidermidis (var. 
A) Kligler (Jour. Infect. Dis., 12, 1913, 
444) which was based on a white culture 
received from Krai under the name 
Micrococcus aurantiacus was apparently 
a white strain of this organism as it grew 
luxurianth^ on ordinary agar. 

12. Micrococcus epidermidis (Winslow 
and Winslow) Hucker. [Staphylococcus 
epidermidis albus Welch, Amer. Jour, of 
Med. Sci., Phila., X. S., 102, 1891, 441; 
Micrococcus epidermidis albus Randolph, 
Jour. Amer. Med. Assoc, 31, 1898, 706; 
Albococcus epidermidis Winslow and Win- 
slow, Syst. Relationships Coccaceae, 
New York, 1908, 201; Staphylococcus 
epidermidis Evans, Jour. Inf. Dis., 15, 
1916, 449; Hucker, X. Y. Agr. Exp. Sta. 
Tech. Bull. 102, 1924, 21.) From Greek, 
epidermis, the outer skin. 

Spheres: 0.5 to 0.6 micron, occurring 
singly, in pairs and in irregular groups. 
Xon-motile. Gram-positive. 

Gelatin stab : White surface growth 
with slow saccate liquefaction. 

Agar colonies: Rather scant, white, 
translucent. 

Broth: Turbid, with white ring and 
sediment. 

Litmus milk: Acid. 

Potato : Limited growth, white. 

Indole not formed. 

Xitrites are produced from nitrates. 

Usually does not utilize XH4H2PO4 
as a source of nitrogen. 

Acid formed from glucose, fructose, 
maltose, lactose and sucrose, but not 
from mannitol, raffinose, salicin or 
inulin. 

Usually fails to hemolyze blood. Xo 
coagulase produced. 

Parasitic rather than pathogenic. 

Aerobic, facultative. 



244 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Optimumtemperature 37°C. 
Source: Originally isolated from small 
stitch abscesses and other skin wounds. 
Habitat : Skin and mucous membranes. 

13. Micrococcus roseus Fliigge. 
(Rosafarbiger Diplococcus, Bumm, Der 
Mikroorganismen der gonorrhoischen 
Schleimhauterkrankungen, 1 Aufl., 1885, 
25; Fliigge, Die Mikroorganismen, 2 
Aufl., 1886, 183; Neisseria roseaTrevisan, 
I generi e le specie delle Batteriacee, 
Milan, 1889, 32; Diplococcus roseus Eisen- 
berg,Bakt.Diag.,3Aufl.,1891,12;MeWs- 
mopedia rosea Dyar, Ann. N. Y. Acad. 
Sci., 8, 1895, 354; Rhodococcus roseus 
Winslow and Rogers, Jour. Inf. Dis., S, 
1906, 545.) From Latin, roseus, rose- 
colored. 

Spheres: 1.0 to 1.5 microns, occurring 
singly and in pairs. Non-motile. Gram- 
variable. 

Gelatin colonies : Rose surface growth 
usually with slow liquefaction. 

Agar colonies: Circular, entire, rose- 
red surface colonies. 

Agar slant: Thick, rose-red, smooth, 
glistening streak. 

Broth : Slightly turbid with rose- 
colored sediment. 

Litmus milk: Unchanged to alkaline, 
usually reddish sediment after 14 days. 

Usually produce nitrites from nitrates. 

Potato: Raised, rose-red, smooth, glis- 
tening. 

Starch not hydrolyzed. 

Acid from glycerol and mannitol. 

Utilizes NH4H2PO4 as a source of 
nitrogen. 

Saprophytic. 

Aerobic. 

Optimum temperature 25°C. 

Source: Dust contamination. 

Habitat: Widespread, as it occurs in 
dust. 

14. Micrococcus cinnabar eus Fliigge. 
(Fliigge, Die Mikroorganismen, 2 Aufl., 
1886, 174; Rhodococcus cinnabareus Wins- 
low and Rogers, Jour. Inf. Dis., 3, 1906, 
545.) From M. L., cinnabar-colored. 



Spheres: 1.0 micron, occurring singly 
and in pairs. Non-motile. Gram-vari- 
able. 

Gelatin colonies : Small, circular, bright 
red, becoming cinnabar red. 

Gelatin stab : Thick, raised, rose to 
cinnabar red growth on surface. No 
liquefaction. White colonies along stab. 

Agar slant: A carmine-red streak. 
Slow growth. 

Broth: Turbid. 

Litmus milk : Slightly alkaline to 
slightly acid. 

Potato : Slowly developing vermillion 
red streak. 

Small amount of acid from test sugars. 

Indole not formed. 

Does not utilize NH4H2PO4 as a source 
of nitrogen. 

Nitrites produced from nitrates. 

Starch not hydrolyzed. 

Saprophytic. 

Aerobic. 

Optimum temperature 25°C. 

Source : Found as contamination of 
cultures. 

Habitat : Usually found as a dust con- 
tamination. 

15. Micrococcus rubens Migula. {Mi- 
crococcus tetragenus ruber Bujwid, in 
Schneider, Arb. bakt. Inst. Karlsruhe, 
1, Heft 2, 1894, 215; Migula, Syst. d. 
Bakt., ^, 1900, 177; Micrococcus ruber and 
Rhodococcus ruber Holland, Jour. Bact., 
6, 1920, 224; Micrococcus roseofulvus 
Hucker, N. Y. S. Agr. Exp. Sta. Tech. 
Bull. 135, 1928, 27; not Micrococcus roseo- 
fulvus Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., ^, 1896, 177 and 439 ; Rhodo- 
coccus roseoftdvus Pribram, Klassifika- 
tion der Schizomyceten, 1933,44). From 
Latin, rubens, ruddy. 

The following description is taken from 
Migula {loc. cit.) and from Hucker {loc. 
cit.) supplemented from unpublished 
notes of the latter. Also see Breed (Jour. 
Bact., 45, 1943,455). 

Spheres: 1.3 to 4.0 microns, average 
size 2.1 microns, occurring in fours and 



FAMILY MICROCOCCACEAE 



245 



in irregular masses, generally not singly 
or in pairs. Non-motile. Gram-nega- 
tive to Gram-variable 

Gelatin colonies : After several days, 
small, pink or flesh-colored, shiny, buty- 
rous, 0.5 to several mm. in diameter. 
Smaller colonies have regular edges ; 
larger colonies have lobate edges. 

Gelatin streak: Thick, shiny, flesh- 
colored to carmine-red growth, generally 
spreading. 

Gelatin stab: Scant, whitish growth 
along line of stab; surface growth flesh- 
red. No liquefaction after several weeks, 
but a slight softening of the medium 
underneath the growth. 

Agar slant: Luxuriant, thick, spread- 
ing, slimy, flesh-colored growth. 

Broth: Bright red, slimy sediment. 
No pellicle. 

Milk: General!}' acid curd followed by 
slight peptonization. 

Nitrites produced from nitrates. 

Acid from glucose, sucrose, mannitol 
and glycerol. No action on lactose or 
starch. 

Pigment soluble in ether, benzol, car- 
bon bisulfide, chloroform and alcohol. 
Not soluble in water (Schneider, loc. 
cit.). 

Saprophytic. 

Grows well at 26" to .37°C. 

Aerobic. 

Source : Original culture isolated by 
Bujwid in Bern, Switzerland and sent to 
Migula at Karlsruhe, Germany. 

Habitat: Unknown. 

16. Micrococcus rhodochrous (Zopf) 
-Migula. (Rhodococcus rhodochrous Zopf, 
Berichte d. deutsch. bot. Gesellsch., 9, 
1891, 22; Migula, Syst. d. Bakt., 3, 1900, 
162.) From Greek, rhodiim, rose; chros, 
color. 

Spheres: 0.5 to 1.0 micron, occurring 
singly. Non-motile. Gram-variable. 

Gelatin colonies: Small, circular, glis- 
tening, raised, entire, dark, reddish- 
brown . 



Gelatin stab: Dark, carmine-red, dry 
surface growth. Slight growth in stab. 
No liquefaction. 

Agar slant : Carmine-red streak, be- 
coming brick-red in color. 

Broth : Thick rose-red pellicle with 
red, flocculent sediment. 

Litmus milk : Slightly alkaline. 

Potato: Carmine-red streak. 

Does not ferment glycerol and man- 
nitol. 

Aerobic. 

Saprophytic. 

Optimum temperature 25°C. 

Habitat : Water. 

17. Micrococcus agilis Ali-Cohen. 
(Ali-Cohen, Cent. f. Bakt., 6, 1889, 36; 
Planosarcina agilis Migula, in Engler 
and Prantl, Die naturl. Pflanzenfam., 1, 
la, 1895, 20; Micrococcus agilis ruber 
Peppier, Cent. f. Bakt., I Abt., 29, 1901, 
352; Planococcus agilis Chester, Man. 
Determ. Bact., 1901, 115; Rhodococcus 
agilis Winslow and Rogers, .Jour. Inf. 
Dis., 3, 1906, 545; Sarcina agilis Ender- 
lein, Sitzber. Gesell. Naturf. Freunde 
Berlin, 1930, 182; not Sarcina agilis 
Matzuschita, Zeit. f. Hyg., 35, 1900, 
496; not Sarcina agilis Saito, Jour. Coll. 
Sci. Imp. Univ. Tokyo, 23, 1908, .) 
From Latin, agilis, agile. 

Spheres, 1.0 micron, occurring singly, 
in pairs and in fours. Motile by means 
of one or two flagella. Gram-variable. 

Gelatin colonies : Small, gray, becoming 
distinctly rose-colored. 

Gelatin stab: Thin, whitish growth in 
stab. On surface thick, rose-red, glis- 
tening growth. Generally no liquefac- 
tion . 

Agar slant: Glistening, dark rose-red, 
lobed, much variation in color. 

Broth: Slightly turbid, with slight, 
rose-colored ring and pink sediment. 

Litmus milk: Slightly acid, pink sedi- 
ment . 

Potato: Slow growth as small, rose- 
colored colonies. 

Loeflier's blood serum: Pink, spread- 



246 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



ing, shiny, abuudaut. Slow liquefaction. 

Indole not formed. 

Nitrites produced (trace). 

Ammonia formed (trace). 

Does not utilize NH4H2PO4 as source 
of nitrogen. 

Acid from glucose, sucrose, inulin, 
glycerol and mannitol. No acid from 
raffinose . 

Aerobic. 

Saprophytic. 

Optimum temperature 25°C. 

Source : Isolated from water. 

Habitat : Water, sea water, on sea fish. 

*18. Micrococcus aerogenes (Schott- 
miiller) Bergey et al. {StapMjlococcus 
aerogenes Schottmliller, Cent. f. Bakt., I 
Abt., Orig., 64, 1912, 270; Bergey et al., 
Manual, 1st ed., 1923, 70; not Micrococcus 
aerogenes Miller, Deutsch. mod. 
Wchnschr., 12, 1886, 119.) From Greek, 
forming air or gas. 

Description according to Prevot, Ann. 
Sci. Nat., Ser. Bot. et Zool., 15, 1933, 
212. 

Spheres: 0.6 to 0.8 micron, occurring 
in clusters, sometimes in pairs or short 
chains. Gram-positive. 

Gelatin: No liquefaction. 

Deep agar colonies: Small, lenticular, 
nearly spherical, yellowish white. Some 
gas bubbles produced, not fetid. 

Blood agar colonies: Very small, gray- 
ish. No true hemolysis, but a narrow 
clear zone is formed. 

Serum agar: Colonies lenticular. Gas 
not fetid. 

Neutral red serum agar : Colonies len- 
ticular. Gas produced. Neutral red 
changed to greenish yellow. 

Glucose broth with blood: Turbid. 
Gas produced. Hydrogen sulfide not 
produced. Slight hemolysis. 

Glucose serum broth: Turbid. Gas 
produced. 

Peptone water with serum : Gas . Indole 
produced. 



Milk: Growth feeble. Neither acid 
nor coagulated. 

Proteins not attacked. 

Glucose and fructose attacked slightly 
by two out of three strains. 

Does not plasmolyse readily. 

Neutral red broth : Changed to yellow- 
ish green. 

Nitrites not produced from nitrates. 

Optimum pH 6.5 to 8.0. 

Optimum temperature 37°C. 

Pathogenic. 

Strict anaerobe. 

Distinctive character: Fermentation 
of glucose and gas production from 
peptones. 

Source : Isolated (Schottmliller) from 
cases of puerperal fever. Three strains 
from infected tonsils studied by Prevot. 

Habitat : Natural cavities, especially 
the tonsils and female genital organs. 

19. Micrococcus asaccharolyticus (Dis- 
taso) comb. nov. {Staphylococcus asac- 
charolyticus Distaso, Cent. f. Bakt., I 
Abt., Orig., 62, 1912, 445.) From Greek, 
not dissolving sugar. 

Description according to Prevot, Ann. 
Sci. Nat., Ser. Bot., 15, 1933, 211. 

Large spheres: 1.0 to 1.2 microns, 
occurring in very large clusters, also in 
pairs and short chains. Gram-positive. 

Gelatin: At 37°C, growth resembles 
tufts of cotton wdiich precipitate. No 
liquefaction. 

Deep agar colonies: Very delicate, 
pin-point, transparent. A few bubbles 
of gas produced. 

Broth: Turbid. Growth settles at 
the bottom of the tube as a sort of vis- 
cous zooglea. Unpleasant odor pro- 
duced. 

Peptone water: Turbid. Indole pro- 
duced. 

Milk: Feebly acidified, but not 
coagulated. 

Egg white not attacked. 

Carbohydrates not attacked. 

Strict anaerobe. 



Anaerobic section revised by Dr. Ivan C. Hall, New York, N. Y. 



FAMILY MICROCOCCACEAE 



247 



Distinctive characters: Large size; 
unpleasant odor; production of indole; 
production of gas. 

Source: Isolated from the large in- 
testine of a man with intestinal in- 
toxication. 

Habitat: Intestine. Not common. 

Note: Weinberg, Nativclle and Prevot 
(Les Microbes Anaerobies, 1937, 1023) 
regard Micrococcus indolicus Christian- 
sen (Ac. Pat. Micr. Scand., 18, 1934, 42) 
as a variety of this species giving it the 
name Staphylococcus asaccharolyticus 
var. indolicus. This variety differs 
from the species by forming opaque 
lens-shaped colonies and by a more 
abundant production of gas from 
peptone. 

20. Micrococcus niger Hall. (Jour. 
Bact., ^0, 1930, 409.) From Latin, niger, 
black. 

Small spheres: 0.6 micron in diameter, 
occurring in irregular masses, occasionallj- 
in pairs. Gram-positive. 

Gelatin: After 5 days a dark sediment 
is produced which gradually gets more; 
and more intensely black. No lique- 
faction. 

Deep agar colonies: Slow growth. At 
first very tiny, colorless, irregularly 
globular, smooth, dense. Small bubbles 
of gas sometimes produced. After sev- 
eral days colonies become brown, then 
black. If exposed to air, colonies fade 
to a dull gray. Medium not discolored. 

Blood agar slant: After 4 or 5 days, 
minute, black colonies, round, smooth, 
glistening, 0.5 mm. in diameter. Non- 
hemolytic. 

Broth : After 4 or 5 days uniform tur- 
bidity and slight production of gas which 
contains H2S. Black sediment. 

Coagulated serum: Minute, brown 
colonies appear on the 8th day. No 
liquefaction. 

Milk: No change. 

Brain medium : Turbid after 4 or 5 days 
at 37°C. Uniform gas production about 
the Gth day. Discoloration of tiie me- 
dium not marked. 



No acid from carbohydrates. Black 
sediment produced. 

Non-pathogenic for guinea-i)igs and 
rabbits. 

Optimum temperature 37°C. No 
growth below 30°C. 

Strict anaerobe. 

Distinctive characters : Formation of a 
water-insoluble, black pigment. Growth 
slow, visible after 2 to 4 days. 

Source: Isolated from urine of an aged 
woman. 

Habitat: Unknown. 

21. Micrococcus grigorofii Prevot. 
(Micrococcus A, Grigoroif, These de 
Geneve, 1905; Prevot, Ann. Sci. Nat., 
Ser. Bot. et Zool., IS, 1933, 219.) Named 
for Grigoroff, who first isolated this 
organism. 

Small spheres : Average size 0.7 micron, 
occurring singly or in irregular masses. 
Gram-positive. 

Gelatin : Colonies appear in four days. 
No liquefaction. 

Deep agar colonies: After three days, 
round, lenticular, yellowish. 

Glucose broth : Turbid after 2 days 
with whitish sediment. Neither gas 
nor fetid odor produced. The medium 
is acidified. 

Milk: Good growth. Acid. Coagu- 
lation. 

Acid from glucose, maltose, lactose, 
fructose and sorbitol. 

One strain slightly pathogenic. 

Optimum temperatvu'e 37°C. 

Strict anaerobe. 

Distinctive characters : This is the 
only anaerobic coccus growing in irregular 
masses that coagulates milk. Lactose is 
fermented. 

Source: Five strains isolated from the 
appendix by Grigoroff. One strain iso- 
lated from an appendix by Prevot. 

Habitat : Human digestive tract. Not 
common. 

22. Micrococcus anaerobius (Hamm) 
comb. 710V. ("Anaerobic staphylococcus, 
Jungano, Compt. rend. Soc. Biol. Paris, 
59, 1907, 707; Sta'phylococcus anaerobius 



248 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Hamm, Die puerperale Wundinfektion, 
Berlin, 1912; not Staphylococcus anaero- 
bius Heurlin, Bakt. Unters. d. Keimge- 
haltes im Genitalkanale d. fiebernden 
Wochnerinnen, Helsingfors, 1910, 120.) 
From Greek, living without air. 

Description according to Pr^vot, Ann. 
Sci. Nat., S^r. Bot., 15, 1933, 209. 

Small spheres: 0.5 to 0.6 micron, 
occurring in masses. Gram-positive. 

Gelatin: No liquefaction. 

Deep agar colonies: Lenticular, thick. 
No gas produced. 

Broth: Turbid, later clearing. Sedi- 
ment. 

Glucose broth: Good growth. Nei- 
ther acid nor gas produced. 

Peptone water: No turbidity. No 
gas. Indole not produced. 

Milk: Neither coagulated nor acidified. 

Coagulated serum not attacked. 

Egg white not attacked. 



Carbohydrates not attacked by the 
strains of Jungano. Acid feebly pro- 
duced from glucose and galactose by 
Prevot's strain. 

Does not plasmolyse. 

Temperature relations: Optimum 36° 
to 38°C. At 22°C growth slow, poor. 
No growth below 22°C. Killed in ten 
minutes at 80°C or in half an hour at 
60°C. 

Optimum pH 6.0 to 8.0. 

Pathogenic for guinea-pigs and 
rabbits. 

Strict anaerobe. 

Distinctive characters: Neutral red 
broth remains unchanged. No gas 
produced. 

Source: First isolated by Jungano 
from a case of cystitis . Found by Pr^vot 
in the pus from a suppurated tonsil. 

Habitat: Urinary tract, urethra, in- 
testine, buccal cavity and conjunctiva. 



Appendix I*: The following genus is organized ona physiological basis. Because of 
this no attempt is made to fit it into the classification outline. A single specieshas 
been described. 

Genus A. Methanococcus Kluyver and van Niel. 
(Cent. f. Bakt., II Abt., 94, 1936, 400.) 
Spherical cells, occurring singly or in masses. Motility not observed. No endo- 
spores formed. Gram-variable. Chemo-heterotrophic, anaerobic, fermenting vari- 
ous organic compounds with the formation of methane. Saprophytes. 
The type species is Methanococcus mazei Barker. 



1. Methanococcus mazei Barker. 
(Pseudosarcina, Maze, Compt. rend. 
Soc. Biol., Paris, 78, 1915, 398; Barker, 
Arch, f . Mikrobiol., 7, 1936, 430.) Named 
for Maze, the French bacteriologist who 
first gave a clearly recognizable descrip- 
tion of this type of methane organism. 

Small spherical cells, occurring singly, 
in large, irregular masses, or in regular 
cysts of various sizes and forms. Non- 
motile. Stains readily with erythrosine. 
Gram-variable. 

Grows on calcium acetate enrichment 
media and ferments the acetate vigor- 
ously. 



Grows slowly on agar containing 2 per 
cent clear mud extract. 

Ferments acetic and butyric acids with 
production of methane in the presence of 
CO2. Ethyl and butyl alcohols not 
attacked. 

Does not utilize organic nitrogen. 

Obligate anaerobe. 

Grows best at 30° to 37°C. 

Sources: Garden soil, black mud con- 
taining HoS, feces of herbivorous ani- 
mals. 

Habitat : One of the most active meth- 
ane-producing organisms found in nature. 



* Appendixes T and II prepared by 
Experiment Station. Geneva, New York 



Prof. Robert S. Breed, New York State 
December, 1943. 



FAMILY MICROCOCCACEAE 



249 



Appendix II: The following genus is recognized by workers in the brewing industry. 
It includes species that present characters intermediate between Micrococcus, Sarcina 
and Sireptococctis . Many students prefer to regard these as species of Micrococcus 
(Hucker, N. Y. State Exper. Sta., Tech. Bui. 102, 1924, 5), of Sarcina (Mace, Traite 
pratique d. Bact., 4th ed., 1901, 460) or of Streptococcus (Shimwell, Sect. 670 in 
Hind, Brewing Science and Practice, New York, 1940). Others (Mees, Thesis, Delft, 
1934) would include in the genus, the species described as Tetracoccus by Orla -Jensen 
(The Lactic Acid Bacteria, Copenhagen, 1919, 76). 

Genus B. Pediococcus Balcke. 
(Wchnschr. f. Brauerei, 1, 1884, 257.) 

Cocci occurring singh^, in pairs and tetrads. Xon-motile. No endospores. Gram- 
positive. Facultative anaerobes under favorable conditions, especially in acid media. 
Nitrites not produced from nitrates. Produce acidification and more or less clouding 
of wort and beer. Saprophytes. 

The tj'pe species is Pediococcus cerevisiae Balcke. 



1. Pediococcus cerevisiae Balcke. 
(Ferment No. 7, Pasteur, Etudes sur la 
biere, Paris, 1876, 4; ySarcina Balcke, 
Wchnschr. f. Brauerei, /, 1884, 183; 
ibid., 1, 1884, 257 ; Merismopedia cerevisiae 
Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 
348; Micrococcus cerevisiae Migula, Syst. 
d. Bakt., 2, 1900, 77; Sarcina cerevisiae 
Mace, Traits Pratique d. Bact., 4th ed.. 
1901, 460.) From Latin, cerevisia, beer. 

Spheres: 1 to 1.3 microns, occurring 
singly, in pairs or tetrads. In acid media 
the latter prevail. Catalase negative. 
Non-motile. Gram-positive. 

No growth in alkaline media. 

Peptone, meat-e.xtract gelatin : White 
becoming yellowish to yellowish brown. 
No liquefaction. 

Wort gelatin with Ca-carbonate : White 
colonies, 2 to 3 mm; carbonate dissolved. 

Meat extract gelatin stab: Growth 
along stab, white raised surface growth. 
No liquefaction. 

Litmus milk: No growth. 

Potato: Scanty growth. 

Acid from glucose, fructose, maltose, 
sucrose. 

Wort and beer : Slight to moderately 
turbid growth, strong development on 
bottom of the flask. Hop sensitive, but 
may develop in heavily hopped beers 
under special conditions. 



Does not utilize urea. 
Nitrites not produced from nitrates. 
Facultative anaerobic. 
Killed at 60°C. in 8 minutes. Optimum 
temperature: 25°C. 
Source : Sarcina-sick beer. 
Habitat: Wort, beer and beer yeast. 

Additional species have been described 
from spoiled wort and beer which vary 
but slightly from the species first named 
and described by Balcke. These are 
listed below together with other species 
that have been placed in the genus. 

Pediococcus acidilactici Lindner. 
(Lindner, Wchnschr. f. Brauerei, 3, No. 
23, 1887, see Cent. f. Bakt., 2, 1887, 342; 
also see Die Sarcina-Organismen der 
Gahrungsgewerbe, Lindner, Inaug. Diss., 
Berlin, 1888, 26, and Cent. f. Bakt., 4, 
1888, 429; Micrococcus pseudocerevisiae 
Migula, Syst. d. Bakt., 2, 1900, 77; Mi- 
crococcus acidi-lactici Chester, Man. 
Determ. Bact., 1901, 88.) From spoiled 
mash. 

Pediococcus albus Lindner. (Die Sar- 
cina-Organismen der Gahrungsgewerbe, 
Lindner, Inaug. Diss., Berlin, 1887, 39; 
see Cent. f. Bakt., 4, 1888, ^29; Micrococ- 
cus pseudosarcina Migula, Sj^st. d. Bakt., 
2, 1900, 92; Micrococcus albus Chester, 
Man. Determ. Bact., 1901, 97.) From 
spoiled beer. 

Pediococcus damnosus Claussen. 



250 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



(Compt. rend. Trav. Labor, de Carlsberg, 
6, 1906, 68; Streptococcus dajnnosus Shim- 
well and Kirkpatrick, Jour. Inst. Brew- 
ing, 45, 1939, 137.) From clear, spoiled 
beer. 

Pediococcus halophilus Mees. (Tetra- 
coccus No. 1, Orla-Jensen, The Lactic 
Acid Bacteria, 1919, 77; Mees, Thesis, 
Delft, 1934, 94.) From anchovy pickle. 

Pediococcus hennebergi Sollied. 
(Ztschr. Spiritusindus., 26, 1903, 491.) 
From spoiled beer. 

Pediococcus kochii Trevisan. (Mikro- 
kokkus in Wundsecreten bei Menschen, 
Koch; Trevisan, I generi e le specie delle 
Batteriacee, Milan, 1889, 28.) 

Pediococcus pentosaceus Mees. (Tetra- 
coccus No. 2, Orla-Jensen, The Lactic 
Acid Bacteria, 1919, 78; Mees, Thesis, 
Delft, 1934, 94.) From yeast. 

Pediococcus perniciosus Claussen (loc. 
cit.). From clouded, spoiled beer. 

Pediococcus sarcinaeformis Rei chard. 
(Ztschr. f. d. ges. Brauwesen, 17, 1894, 
257.) From spoiled beer. 

Pediococcus urinae equi Mees. (Pfer- 
deurinsarcina, von Huth, Alg. Ztg. f. 
Bierber. u. Malzfabr., 185, 968 and 981, 
1885; ibid., 1886, 141; Mees, Thesis, 
Delft, 1934, 95.) From horse urine. 

Pediococcus violaceus (Klitzing) Trevi- 
san. {Merismopedia violacea Klitzing; 
Trevisan, I generi c le specie delle Bat- 
teriacee, Milan, 1889, 28.) 

Pedioplana haeckeli Wolff. (Cent. f. 
Bakt., II Abt., 18, 1907, 9.) Motile. 
From rotting beets. Placed in a new 
genus Pedioplana Wolff {loc. cit., 9). 

Streptococcus damnosus var. mucosus 
Shimwell. (Shimwell, Sect. 670, Hind, 
Brewing Science and Practice, New 
York, 1940.) From ropy beer. 

Appendix III*. The following species 
have been found in the literature and are 
listed here chiefly for their historical 
interest. Many are incompletely de- 
scribed, while many others are identical 



with previously described species. See 
Monographs by Winslow and Winslow, 
Sj'stematic Relationships of the Cocca- 
ceae, 1908 and Hucker, N. Y. State Exper. 
Sta., Tech. Buls. Nos. 99-103. Refer- 
ences are to Tech. Bui. 102. 

Ascococcus cantabridgensis Hankin. 
(Quoted from Lehmann and Neumann, 
Bakt. Diag., 2 Aufl., 2, 1899, 165.) Mi- 
gula (Syst. d. Bakt., 2, 1900, 195) reports 
he is unable to find further reference to 
this organism and we likewise are unable 
to trace it. From the human mouth. 

Ascococcus gangrenosus Bevan. (Med. 
News, No. 1003, 1892, 375; Abst. in Cent, 
f. Bakt., 13, 1893, 796.) From a gan- 
grenous foot. 

Ascococcus vibrans van Tieghem. (Bui. 
Soc. Bot. France, 27, 1880, 150.) From 
water. 

Aurococcus tropicus Chalmers and 
O'Farrell. (1913, quoted from Castel- 
lani and Chalmers, Man. Trop. Med., 
3rd ed., 1919, 931.) Found in Ceylon in 
granulating ulcers of skin. 

Coccus carduus Heurlin. (Bakt. Un- 
ters. d. Keimgehaltes im Genitalkanale 
d. fiebernden Wochnerinnen. Helsing- 
fors, 1910, 136.) Anaerobic. From 
genital canal. 

Coccus caudatus Heurlin {loc. cit., 84). 
From genital canal. 

Coccus vaginalis Heurlin {loc. cit., 79). 
From genital canal. 

Galactococcus albus Guillebeau. (Land- 
wirtsch. Jahrb. d. Schweiz, 4, 1892, 27; 
Abst. in Cent. f. Bakt., 12, 1892, 101.) 
From milk from an inflamed udder. 

Galactococcxis fulvus Guillebeau {loc. 
cit.). From milk from an inflamed udder. 

Galactococcus versicolor Guillebeau 
{loc. cit.). From milk from an inflamed 
udder. 

Gyrococcus flaccidifex Glaser and Chap- 
man. (Science, 36, 1912, 219.) Isolated 
from the gypsy moth, Porthetria dispar. 

Jodococcus vaginatus Miller. (Miller, 



* Prepared for Prof. G. J. Hucker by Mrs. Eleanore Heist Clise, New York State 
Experiment Station, Geneva, New York, March, 1943. . 



FAMILY MICROCOCCACEAE 



251 



Mikroorganismen der Mundhohle, 1889, 
54; Bacterium iogenum Baumgartner, 
Ergebnisse d. ges. Zahnheilk., Heft 2, 
1910, 729; Abst. in Cent. f. Bakt., I Abt., 
Ref., 48, 1911, 621.) From the oral 
cavity. 

Merismopedia auranliaca Maggiora. 
(Giorn. Soc. Ital. d'Igiene, 11, 1889, 354; 
Abst. in Cent. f. Bakt., 8, 1890, 13.) 
From the normal skin of the human foot. 

Micrococcu!^ achrous Migula. (No. 16, 
Lembke, Arch. f. Hyg., 26, 1896, 310; 
Migula, Syst. d. Bakt., 2, 1900, 201.) 
From feces. Winslow and Winslow (Sys- 
tematic Relationships of the Coccaceae, 
1908, 224) state that this species is appar- 
ently a synonym of Micrococcus candi- 
cans Fliigge. 

Micrococcus acidi lactici Marpmann. 
(Erganzungsheft d. Cent, f . allg. Gesund- 
heitspflege, 2, 1886, 22.) Found in fresh 
milk. 

Micrococcus acidovorax MuUer-Thur- 
gau and Osterwalder. (Cent. f. Bakt., 
II Abt., 86, 1913, 236.) From wine. 
Hucker {loc. cit., 6) considers this a 
synonym of Micrococcus luteus Cohn or 
Micrococcus varians Migula. 

I\[icrococcus acne Hollaml. (Jour. 
Bact., S, 1920, 223; Staphylococcus acne 
Holland, ibid., 225; see Micrococcus cittis 
communis Sabouraud.) 

Micrococcus (Staphylococcus) acridi- 
cida Kufferath. (Ann. de CJembloux, 
27, 1921, 253.) Isolated from diseased 
locusts from Greece. Resembles Micro- 
coccus aureus Zopf. 

Micrococcus aerius Chester. (Xo. 49, 
Conn, Storrs Agr. Exp. Sta. 7th Ann. 
Rept., 1895, 81; Chester, Man. Determ. 
Bact., 1901, 104.) From dust. Hucker 
{loc. cit., 12) states that this species 
appears to be identical with Micrococcus 
aureus Zopf. 

Micrococcus aerogenes Miller. (Miller, 
Deutsche med. Wchnschr., 12, 1886, 119; 
not Micrococcus aerogenes Bergey et al., 
Manual, 1st ed., 1923, 70.) P>om the 
alimentary canal. 



Micrococcus agilis albus Catterina. 
(Cent. f. Bakt., I Abt., Orig., 34, 1903, 
108.) Found in septicemia of rabbits. 
Motile with one or two flagella. 

Micrococcus alhatus Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 479.) 
From the intestine of a woodpecker 
{Picus major). Winslow and Winslow 
(Systematic Relationships of the Coc- 
caceae, 1908, 199) state that this species 
appears to be a synonym of M icrococcus 
albus Schroeter; while Hucker (N. Y. 
Agr. Exper. Sta., Tech. Bull. 102, 19) 
regards it as a synonym of Micrococcus 
freudenreichii Guillebeau or M icrococcus 
ureae Cohn. 

Micrococcus albescens Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
76.) From cheese. Winslow and Wins- 
low (loc. cit., 199) state that this species 
appears to be a synonym of Micrococcus 
albus Schroeter; while Hucker (loc. cit., 
19) regards it as a synonym of Micrococ- 
cus freudenreichii Guillebeau or of Mi- 
crococcus ureae Cohn. 

Micrococcus albidus Losski. (Losski, 
Inaug. Diss., Dorpat, 1893, 55; not 
Micrococcus albidus Henrici, see Micro- 
coccus subniveus below; not Micrococcus 
albidus Roze, Compt. rend. Acad. Sci. 
Paris, 122, 1896, 750.) From soil. 
Hucker (loc. cit., 19) regards this species 
as a synonym of Micrococcus freuden- 
reichii Guillebeau or Micrococcus ttreae 
Cohn. 

Micrococcus albocereus Migula. (Sta- 
phylococcus cereus albus Passet, Unter- 
such. ii. d. Aetiol. d. eiterigen Phlegmone 
d. Menschen, Berlin, 1885, 53, and Fort- 
schr. d. Med., 3, 1885; Micrococcus cereus 
albus Fltigge, Die Mikroorganismen, 2 
Aufl., 1886, 182; Staphylococcus cereus 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 32; Migula, Syst. 
d. Bakt., 2, 1900, 56; Staphylococcus 
cereus-albus Holland, Jour. Bact., 5, 1920, 
225.) From human pus, also from water. 
Winslow and Winslow (Systematic Rela- 
tionships of the Coccaceae, 1908, 205) 
consider this a synonym of M icrococcus 



252 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



candidus Cohn or of Gaffkya tetragena 
Trevisan. 

Micrococcxis albus Frankland and 
Frankland. (Phil. Trans. Roy. Soc, Lon- 
don, 178, B, 1888, 264; not Micrococcus 
albus Matzuschita, Cent. f. Bakt., I 
Abt., 29, 1901, 382; not Micrococcus 
albus Buchanan, Veterinary Bacteri- 
ology, 1911, 196; not Micrococcus albus 
Mace, Traits Pratique de Bact., 6th 
^d., 1912, 605.) From air. Resembles 
Micrococcus candicans. 

Micrococcus albus II Maggiora. 
(Cent. f. Bakt., 8, 1890, 13.) See Micro- 
coccus opalescens De Toni and Trevisan. 
From the skin of the human foot. 

Micrococcus ampins Migula. (Grau- 
weisser Diplococcus, Bumm, Der Mikro- 
org. d. gonorrh. Schleimhauterkrank., 
1 Aufl., 1885, 17; Micrococcus albicans 
ampins Fliigge, Die Mikroorganismen, 2 
Aufl., 1886, 183; Neisseria albicans 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 32; Diplococcus 
albicans amplus Eisenberg, Bakt. Diag., 
3 Aufl., 1891, 24; Migula, Syst. d. Bakt., 
2, 1900, 118 ; Micrococcus albicans Chester, 
Man. Determ. Bact., 1901, 80.) From 
vaginal secretions. Hucker (loc. cit., 
15) considers this species identical with 
Micrococcus albus Schroeter. 

Micrococcus ampullaceus Kern. (Arb. 
bakt. Inst. Karlsruhe, ^ , Heft 4, 1897, 477.) 
From the intestine of a dove (Columba 
oenas). Hucker (loc. cit., 19) considers 
this a synonym of Micrococcus freuden- 
reichii Guillebeau or Micrococcus ureae 
Cohn. 

Micrococcus annulatus Kern {loc. cit., 
490). From the stomach contents of the 
hedge sparrow (Passer montanus) and 
the intestine of the rock dove {Columba 
livia). Winslow and Winslow (Syste- 
matic Relationships of the Coccaceae, 
1908, 216) consider this species a synonym 
of Micrococcus flavus Lehmann and Neu- 
mann. 

Micrococcus aquatilis Bolton. (Zschr. 
f. Hyg., /, 1886, 94; not Micrococcus 
aquatilis Chester, see below.) From 



water. Winslow and Winslow {loc. cit., 
224) state that this species is apparently 
a synonym of Micrococcus candicans 
Flugge. 

Micrococcus aquatilis Chester. {Mi- 
crococcus aquatilis invisibilis Vaughan, 
Amer. Jour. Med. Sci., /04, 1892, 183; 
Chester, Man. Determ. Bact., 1901, 88.) 
From water. Winslow and Winslow {loc. 
cit., 224) state that this species is ap- 
parently a synonym of Micrococcus candi- 
cans Flugge. 

Micrococcus aquatilis albus. (Quoted 
from Toporoff, Protok. d. Kaiserl. kau- 
kasisch Mediz. Gesellsch., 1892, No. 21; 
Abst. in Cent. f. Bakt., 13, 1893, 487.) 
From water. 

Micrococcus aquatilis flavus. (Quoted 
from Toporoff, loc. cit.) From water. 

Micrococcus aqueus Migula. (No. 25, 
Lembke, Arch. f. Hyg., 26, 1896, 317; 
Migula, Syst. d. Bakt., 2, 1900, 204.) 
From feces. Winslow and Winslow {loc. 
cit., 184) state that this species is appar- 
ently a synonym of Micrococcus aureus 
Zopf, while Hucker {loc. cit., 15) regards 
this as a synonym of Micrococcus albus 
Schroeter. 

Micrococcus arbor escens lactis Conn. 
(Conn, Storrs Agr. Exp. Sta. 12th Ann. 
Rept., 1900, 46; Micrococciis lactis ar- 
borescens Conn, Esten and Stocking, 
Storrs Agr. Exp. Sta. 18th Ann. Rept., 
1907, 110.) From milk. Hucker {loc. 
cit., 21) regards this as a synonym of 
Micrococcus candidus Cohn or Micrococ- 
cus epidermidis Hucker. 

Micrococciis argenteus Migula. (No. 
27, Lembke, Arch. f. Hyg., 26, 1896, 317; 
Migula, Syst. d. Bakt., 2, 1900, 206.) 
From feces. Winslow and Winslow {loc. 
cit., 184) state that this species is appar- 
ently a synonym of Micrococcus aureus 
Zopf, while Hucker {loc. cit., 10) con- 
siders it a synonj'm of Micrococcus con- 
glomcratus Migula. 

Micrococcus ascoformans J ohne . {Zoo- 
gloea pulmonis equi Bollinger, Arch. f. 
path. Anat., 49, 1870, 583; Discomyces 
equi Rivolta, Giorn. di Anat. e Fisiolog., 



FAMILY MICROCOCCACEAE 



253 



10, 1884; Johne, Ber. u. d. Veterin. im 
Konigr. Sachsen, Jahr 1885, 47; Ascococ- 
cus johnei Cohn in letter to Johne, 
Deutsche Ztschr. f. Thiermed., 12, 1886, 
210; Micrococcus botryogenus Rabe, Deut. 
Ztschr. f. Thiermed., 12, 1886, 137; 
Botryomyces equi Bollinger, Deut. 
Ztschr. f. Thiermed., 13, 1887, 176; 
Botryococcus ascoformans Kitt, Cent. f. 
Bakt., 3, 1888, 247; BoUingera equi 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 26; Staphylococcus 
ascoformans Ford, Textb. of Bact., 1927, 
424; Staphylococcus aureus var. equi 
Hauduroy et al.. Diet. d. Bact. Path., 
1937, 504.) Causes botryomycosis in 
horses. Lehmann and Neumann (Bakt. 
Diag., 7 Aufl., 2, 1927, 291) consider this 
a form of Micrococcus aureus Zopf ; while 
Hucker {loc. cit., 15) regards this as a 
form of Micrococcus albus Schroeter. 

Micrococcus ascoformis Fermi. (Arch, 
f. Hyg., 10, 1890, 10.) Presumably in- 
tended for Micrococcus ascoformans Johne. 

Micrococcus asper Migula. (Seibert, 
Inaug. Diss., Wtirzburg, 1894, 12; Migula, 
Syst. d. Bakt., 2, 1900, 82.) From a 
hairbrush. Winslow and Winslow (loc. 
cit., 205) consider this species to be a 
synonym of Micrococcus candidus Cohn 
or of Gaffkya tetragena Trevisan. 

Micrococcus aurantiacus-sorghi Bruyn- 
ing. (Arch. Neer. Sci. Exact, et Nat., 
/, 1898, 297; Streptococcus aurantiacus 
Chester, Man. Determ. Bact., 1901, 69.) 
From sorghum. 

Micrococcus aureus Chester. (Micro- 
coccus cremoides aureus Dyar, Ann. N. Y. 
Acad. Sci., 8, 1895, 349; Chester, Manual 
Determ. Bact., 1901, 99.) From dust. 
Regarded by Dyar as a varietal form of 
Micrococcus cremoides Zimmermann. 
Winslow and Winslow (loc. cit., 184) 
consider this species a synonym of Mi- 
crococcus aureus Zopf. 

Micrococcus aureus lactis Conn. (Storrs 
Agr. Exp. Sta. 12th Ann. Rept., 1900, 
36.) From milk. This seems to be 
identical with Micrococcus lactis aureus 
A, Conn, Esten and Stocking, Storrs 



Agr. Exper. Sta. 18th Ann. Rept., 1907, 
119. Hucker (loc. cit., 9) regards this 
species as identical in part with Micrococ- 
cus jlavus Lehmann and Neumann and 
with Micrococcus conglotneratus Migula. 

Micrococcus (Sarcina) baccatus Migula. 
(No. 18, Lembke, Arch. f. Hyg., ^6, 1896, 
311; Migula, Syst. d. Bakt., 2, 1900, 202.) 
From feces. Winslow and Winslow (loc. 
cit., 232) state that this is a yellow, gela- 
tin-liquefying sarcina, apparently a syn- 
onym of Sarcina flava De Bary . Hucker 
(loc. cit., 10) considers this a synonym of 
Micrococcus conglomeratus Migula. 

Micrococcus badius Lehmann and Neu- 
mann. (Bakt. Diag., 1 Aufl., 2, 1896, 
163.) Received from the Krai collection 
as Sarcina lutea. Winslow and Winslow 
(loc. cit., 216) consider this a synonym 
of Micrococcus flavus Lehmann and Neu- 
mann. 

Micrococcus baregensis purpureus Ro- 
bine and Hauduroy. (Compt. rend. Soc. 
Biol., Paris, 98, 1928, 25.) From hot 
sulfur springs at Bareges. 

Micrococcus beigelii (Rabenhorst) Mi- 
gula. (Pleurococcus beigelii Kiichen- 
meister and Rabenhorst, Hedwigia, 1867, 
No. 4; Sclerotium beigelianum Hallier, 
1868; Zoogloea beigeliana Eberth, 1873; 
Hyalococcus beigelii Schroeter, Kryptog.- 
Flora V. Schlesien, 3, 1, 1886, 152; 
Chlamydatomus beigelii Trevisan, Rendi- 
conti Reale Inst. Lombardo de Sci. e 
Lett., Ser. II, 12, 1879, 22; Migula, Syst. 
d. Bakt., 2, 1900, 193; Trichosporum 
beigelii Vuillemin, 1901.) From human 
hair. 

Micrococcus beri-beri Pekelharing. 
(Pekelharing, Weekblad v. h. Ned. 
Tijdschr. v. Geneesk., No. 25; also 
Pekelharing and Winkler, Deut. med. 
Wchnschr., No. 39, 1887, 845; Neisseria 
loinkleri Trevisan, I generi e le specie 
delle Batteriacee, Milan, 1889, 32.) Con- 
sidered the cause of beri-beri by Pekel- 
haring. Winslow and Winslow (loc. cit., 
184) state that this is apparently a 
synonym of Micrococcus aureus Zopf; 
while Hucker (loc. cit., 11) considers 



254 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



this a synonym of Micrococcus cilreus 
Migula. 

Micrococcus hicolor Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 485.) 
From the intestine of a dove {Columba 
oenas). Hucker (loc. cit., 21) considers 
this a synonym of Micrococcus candidus 
Cohn or of Micrococcus epidermidis 
Hucker. 

Micrococcus billrothii (Cohn) Migula. 
(Ascococcus billrothii Cohn, Beitr. z. 
Biol. d. Pflanzen, 1, Heft 3, 1875, 151; 
Migula, Syst. d. Bakt., 2, 1900, 195.) 
Found in putrefying meat infusion. 

Micrococcus biskra Heydenreich. 
(Heydenreich, Ausgabe d. Haupt Med.- 
Verhalt., St. Petersburg, 1888; see Cent, 
f. Bakt., 5, 1889, 163; Staphylococcus 
biskrae Trevisan, I generi e le specie delle 
Batteriacee, Milan, 1889, 32; Micrococ- 
cus heydenreichii Chester, Man. Determ. 
Bact., 1901, 91.) Found in ulcers in an 
Oriental skin disease. Winslow and 
Winslow (loc. cit., 184) state that this is 
apparently a synonym of Micrococcus 
aureus Zopf; while Hucker (loc. cit., 
11) considers it a synonym of Micrococ- 
cus cilreus Migula. 

Micrococcus boleti Passerini. (Erbar. 
crittogam. Italiano, II ser.. No. 1199; 
quoted from Trevisan, I generi c le 
specie delle Batteriacee, Milan, 1889, 
34.) Saprophytic on a fungus (Boletus 
edulis). 

Micrococcus bombycis (Naegeli) Cohn. 
(Panhistophyton ovatum Lebert, Jahres- 
ber. i'l. d. Wirksamkeit d. Vereins z. 
Beford. d. Seidenbaues f. Brandenburg 
im Jahre 1856-57, 28; and Berliner En- 
tomolog. Ztschr., 1858; Nosema bombycis 
Naegeli, Botan. Sect. d. 33 Versammlg. 
d. Naturf. u. Aerzte in Bonn, 1857, 160; 
and Botan. Zeitg., 1857, 760; Microzyma 
bombycis Bechamp, Compt. rend. Acad. 
Sci., Paris, 6^, 1867, 1045; Cohn, Beitr. 
z. Biol. d. Pflanzen, 1, Heft 2, 1872, 165; 
Micrococcus ovatus Winter, in Raben- 
horst, Krypt. Flora v. Deutschl., Oes- 
terr. u. d. Schweiz, 2 Aufl., 1, 1884, 47; 
Streptococcus bombycis Zopf, Die Spalt- 



pilze, 2 Aufl., 1884, 52.) Found in the 
blood and organs of diseased silkworms 
(Bombyx mori). 

Micrococcus boreus Issatchenko. (Re- 
cherches sur les Microbes de I'Ocedn 
Glacial Arctique, Petrograd, 1914, 144.) 
From sea water. 

Micrococcus bovinus Migula. (Micro- 
coccus der Lungenseuche der Rinder, 
Poels and Nolen, Fortschr. d. Med., 
1886, 217; Migula, Syst. d. Bakt., 2, 
1900, 67.) From the lungs of diseased 
cattle. Hucker {loc. cit., 22) regards 
this a synonym of Micrococcus candidus 
Cohn or of Micrococcus epidermidis 
Hucker. 

Micrococcus bovis Migula. (Hemato- 
coccus, Babes, Compt. rend. Acad. Sci., 
Paris, 107, 1888, 692 and 110, 1890, 800 
and 975; also see Arch. f. path. Anat., 
115, 1889; Neisseria babesi Trevisan, I 
generi e le specie delle Batteriacee, 
Milan, 1889, 32; Haematococcus bovis 
Eisenberg, Bakt. Diag., 3 Aufl., 1891, 
271 ; Migula, Syst. d. Bakt., 2, 1900, 85.) 
From the blood and organs of cattle. 

Micrococcus burchardti Trevisan. 
(Coccus bei keratitis phlyctaenulosa, 
Burchardt, Cent. f. Bakt., 1, 1887, 392; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 33.) Pathogenic. 
From the cornea of a rabbit. 

Micrococcus butyri (v. Klecki) Migula. 
(Diplococcus butyri von Klecki, Cent, 
f. Bakt., 16, 1894, 358; Migula, Syst. d. 
Bakt ., ^, 1900, 216. ) From rancid butter. 
Winslow and Winslow (loc. cit., 220) con- 
sider this a synonym of Micrococcus 
luteus Cohn. 

Micrococcus butyrlfluorescens Teichert. 
(Inaug. Diss., Jena, 1904; Abst. in Cent, 
f. Bakt., II Abt., IS, 1904, 561.) From 
milk. Exhibits a green fluorescence. 

Micrococcus butyricus (von Klecki) 
Migula. (Tetracoccus butyricus von 
Klecki, Cent. f. Bakt., 15, 1894, 360; 
Migula, Syst. d. Bakt., 2, 1900, 216.) 
From rancid butter. Winslow and Win- 
slow (loc. cit., 220) consider this a syn- 
onym of Micrococcus luteus Cohn. 



FAMILY MICROCOCCACEAE 



255 



Micrococcus calco-aceticus Beijerinck. 
(Proc. Sect. Sci., Kon. Akad. v. Weten- 
schappen, 13, 1911, 1066; Abst. in Cent, 
f. Bakt., II Abt., 31, 1912, 290.) Occurs 
commonly in soils. 

Micrococcus campeneus Orme. (Jour. 
Trop. Med. and Hyg., 11, 1908, No. 10, 
May 15; Abst. in Cent. f. Bakt., I Abt., 
Ref., 43, 1909, 299.) 

Micrococcus candicans Fltigge. (Die 
Mikroorganismen, 2 Aufl., 1886, 173; 
Albococcus candicans Winslow and 
Rogers, Jour. Inf. Dis., 3, 1906, 544; 
Staphylococcus candicans Holland, Jour. 
Bact., S, 1920, 225.) From air, water 
and milk. Hucker (loc. cit., 22) regards 
this a synonym of Micrococcus candidus 
Cohn or of Micrococcus epidermidis 
Hucker. For a description of this spe- 
cies, see Bergey et al., Manual, 5th ed., 
1939, 255. 

Micrococcus canescens Migula. (Mi- 
crococcus No. 4, Adametz, Landwirtsch. 
Jahrb., 18, 1889, 240; Migula, Syst. d. 
Bakt., 2, 1900, 51; Albococcus cariescens 
Winslow and Rogers, Jour. Inf. Dis., 8, 
1906, 544; Staphylococcus canescens Hol- 
land, Jour. Bact., 5, 1920, 225.) From 
Emmenthal cheese. Winslow and Win- 
slow {loc. cit., 224) state that this is 
apparently a synonym of Micrococcus 
candicans Fliigge. 

Micrococcus capillorum (Buhl) Tre- 
visan. (Zoogloea capillorum Buhl, 
Ztschr. f. ration. Med., II Reihe, U, 18-, 
356; Palmella capillorum Ktihn, Abhandl. 
d. Naturf. Ges. zu Halle, 9, Heft 1, 18-, 
62; Palmellina capillorum Rabenhorst, 
Flor. Eur. Alg., 3, 1856 (?), 35;Trevisan, 
I generi e le specie delle Batteriacee, 
Milan, 1889, 33.) From the skin. Con- 
sidered pathogenic. 

Micrococcus capsaformans Jamieson and 
Edington. (Brit. Med. Jour., 1, June 11, 
1887, 1262; Micrococcus capriformis (sic), 
Abst. in Cent. f. Bakt., 2, 1887, 223.) 
From the scales and blood of scarlet 
fever patients. Not pathogenic. 

Micrococcus carbo Renault. (Compt. 
rend. Acad. Sci., Paris, 123, 1896, 935.) 



Micrococcus carneus Zimmermann. 
(Roter Coccus, Maschek. Bakt. Unter- 
such d. Leitmeritz. Trinkwassers, No. 
5, 1887, 60; Zimmermann, Die Bakt. 
unserer Trink- u. Nutzwasser, Chem- 
nitz, I Reihe, 1890, 78.) From water. 
Hucker {loc. cit., 25 and 26) regards 
this species as identical with Micrococcus 
roseus Fliigge or Micrococcus cinna- 
bar eus Fliigge. 

Micrococcus' carnicolor Frankland and 
Frankland. (Phil. Trans. Roy. Soc. 
London, 178, B, 1888, 263; not Micrococ- 
cus carnicolor Kern, see Micrococcus 
subcarneus below.) From air. Hucker 
(loc. cit., 25) states that this species 
may be identical with Micrococcus 
roseus Fliigge. 

Micrococcus carniphilus Wilhelmy. 
(Arb. bakt. Inst. Karlsruhe, 3, 1903, 10.) 
From a meat extract. 

Micrococcus casei amari edamicus 
Raamot. (Inaug.Diss.,K6nigsberg, 1906; 
Abst. in Cent. f. Bakt., II Abt., 18, 
1907, 348.) From pasteurized skim milk. 

Micrococcus castellanii Chalmers and 
O'Farrell. (Ann. Trop. Med. and Para- 
si tol., 7, 1913, 528; Rhodococcus castellanii 
Castellani and Chalmers, Man. Trop. 
Med., 3rd ed., 1919, 2102.) Found in the 
red variety of trichomycosis axillaris, a 
tropical disease. 

Micrococcus cartharinensis Issat- 
chenko. (Recherches sur les Microbes 
de I'Oce^n Glacial Artique, Petrograd, 
1914, 148.) From sea water. 

Micrococcus cellaris (Schroeter) Mi- 
gula. {Leucocystis cellaris Schroeter, 
Kryptog. Flora v. Schlesien, 3, 1886, 152; 
Migula, Syst. d. Bakt., 2, 1900, 195.) 
From a coating on the walls of damp 
cellars and mines. 

Micrococcus centropunctatus Issat- 
chenko. (Recherches sur les Microbes 
de rOcedn Glacial Arctique, Petrograd, 
1914, 146.) From sea water. 

Micrococcus cerasinus ^ligula. (]Mi- 
crococcus aus roter Milch, Keferstein; 
Cent. f. Bakt., I Abt., 21, 1897, 177; .1/z- 
crococcus cerasinus lactis Helm, Lehrb. 



256 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



d. Bakt., 2 Aufl., 1898, 299; Migula, 
Syst. d. Bakt., £, 1900, 170; not Micro- 
coccus cerasinus Lehmann and Neumann, 
Bakt. Diag., 1 AuH., 2, 1896, 179; Micro- 
coccus kefersteinii Chester, Man. De- 
term. Bact., 1901, 107.) From red milk. 
Hucker (loc. cit., 26) regards this species 
as identical with Micrococcus cinna- 
bareus Fliigge. 

Micrococcus cereus Migula. (Staphy- 
lococcus cereus flavus Passat, Unter- 
suchungen liber die Aetiologie der eiteri- 
gen Phlegmone des Menschen, 1885, 53; 
Micrococcus cereus flavus Fliigge, Die 
Mikroorganismen, 2 Aufl., 1886, 182; 
Staphylococcus passeti Trevisan, I generi 
e le specie delle Batteriacee, Milan, 
1889, 32; Migula, Syst. d. Bakt., 2, 1900, 
126; Staphylococcus cereus-flavus Hol- 
land, Jour. Bact., 5, 1920, 225.) From 
pus. Winslovv and Winslow {loc. cit., 
220) consider this species identical with 
Micrococcus luteus Migula. For a de- 
scription of this organism, see Bergey 
et al., Manual, 5th ed., 1939, 241. 

Micrococcus cereus aureus Dyar. 
(Ann. N. Y. Acad. Sci., 8, 1895, 347.) 
Obtained as Staphylococcus cereus aureus 
from Krai's laboratory; also found in air. 

Micrococcus cerinus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
84.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 216) consider this a 
synonym of Micrococcus flavus Trevisan. 

Micrococcus chersonesia Corbet. 
(Quart. Jour. Rubber Research Inst., 
Malaya, 2, 1930, 150.) From the latex 
of the rubber tree {Hevea brasiliensis) . 
For a description of this species, see 
Bergey et al.. Manual, 5th ed., 1939, 
258. 

Micrococcus chinicus Enimerling and 
Abderhalden. (Cent. f. Bakt., II Abt., 
10, 1902, 337.) Putrefying meat. 

Micrococcus chlorinus Cohn. (Griin- 
gelber Coccus, Maschek, Jahresber. d. 
Kom.- Oberrealschule zu Leitmeritz, 
1887, 66; Cohn, Beitr. z. Biol. d. Pflanzen, 
1, Heft 2, 1872, 155.) From water and 
dust. Hucker {loc. cit., 10) considers 



this a synonym of Micrococcus conglom- 
eratus Migula. 

Micrococcus chromidrogenus citreus 
Trommsdorff. (Miinch. Med. Wochn- 
schr., 1904, No. 29, 1286; Abst. in Cent, 
f. Bakt., I Abt., Ref., 37, 1905, 60.) 
Isolated from a case of chromidrosis of 
the axilla. 

Micrococcus chromidrogenus ruber 
Trommsdorff {loc. cit.). Isolated from 
a case of chromidrosis of the axilla. 

Micrococcus chromoflavus Huss. 
(Cent. f. Bakt., II Abt., 19, 1907, 520.) 
From cheese 

Micrococcus chryseus Frankland and 
Frankland. (Phil. Trans. Roy. Soc. 
London, 178, B, 1888, 268.) From dust. 
Winslow and Winslow {loc. cit., 184) 
state that this species is apparently a 
synonym of Micrococcus aureus Zopf. 

Micrococcus cinnabarinus Zimmer- 
mann. (Die Bakt. unserer Trink- u. 
Nutzwasser, Chemnitz, I Reihe, 1890, 
76.) From water. Hucker (Zoc. czi., 26) 
regards this species as identical with 
Micrococcus cinnabareus Fliigge. 

Micrococcus cirrhiformis Migula. 
(Ranken Coccus, Maschek, Jahresber. 
(1. Kom.- Oberrealschule in Leitmeritz, 
1887, 66; Migula, Syst. d. Bakt., 2, 1900, 
53.) From water. Hucker {loc. cit., 22) 
considers this a synonym of Micrococcus 
candidus Cohn or of Micrococcus epider- 
midis Hucker. 

Micrococcus citreus I and //, Maggiora. 
(Giorn. Soc. Ital. d'Igiene, 11, 1889, 354; 
Abst. in Cent. f. Bakt., 8, 1890, 13; not 
Micrococcus citreus Eisenberg, Bakt. 
Diag., 3 Aufl., 1891, 36; not Micrococcus 
citreus Migula, Syst. d. Bakt., 2, 1900, 
147; not Micrococcus citreus Winslow and 
Winslow, Systematic Relationships of 
the Coccaceae, 1908, 218.) From the 
normal skin of the foot. 

Micrococcus citreus granulatus Freund. 
(Inaug. Diss., Erlangen, 1893, 27; Abst. 
in Cent. f. Bakt., 16, 1894, 641 ; Micrococ- 
cus granulatus Bazarewski, Cent. f. 
Bakt., II Abt., 15, 1905, 7; not Micrococ- 
cus granulatus Weiss, Arb. bakt. Inst. 



FAMILY MICROCOCCACEAE 



257 



Karlsruhe, 2, Heft 3, 1902, 197.) From 
the oral cavity. Hucker Q.oc. cit., 9) 
regards this as a synonym of Micrococcus 
flavus Trevisan. 

Micrococcus citreus lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900,. 40.) From milk. Hucker {loc. 
cit., 10) considers this a synonym of 
Micrococcus conglomeratus ]Migula. 

Micrococcus citreus rigensis Bazarew- 
ski. (Cent. f. Bakt., II Abt., 15, 1905, 
5.) From dust. 

Micrococcus citrinus Migula. (Diplo- 
coccus citreus liquefaciens Unna and 
Tommasoli, Monats. f. prakt. Dermatol - 
ogie, 9, 1889, 56; Migula, Syst. d. Bakt., 
2, 1900, 150; Micrococcus tommasoli Ches- 
ter, Man. Determ. Bact., 1901, 101; Mi- 
crococcus citreus liquefaciens Winslow 
and Winslow, Systematic Relationships 
of the Coccaceae, 1908, 216.) From hu- 
man skin in a case of eczema. Winslow 
and Winslow {loc. cit., 216) consider this 
a synonym of Micrococcus flavus Trevi- 
san. 

Micrococcus claviformis von Besser. 
(Beitr. z. path. Anat. u. z. allgem. Path., 
6, 1889, 340; see Cent. f. Bakt., 7, 1890, 
152.) Found once in nasal secretions. 
Micrococcus coccineus Migula. (Mi- 
crococcus No. VI, Adametz,Landwirtsch. 
Jahrb., 18, 1889, 242; Migula, Syst. d. 
Bakt., 2, 1900, 174.) From Emmenthal 
cheese. Hucker (loc. cit., 26) regards 
this species identical with Micrococcus 
cinnabareus Fltigge. 

Micrococcus coli brevis Lehmann. 
(Lehmann, Inaug. Diss., Miinchen, 
1903; Abst. in Cent. f. Bakt., I Abt., 
Ref., 36, 1905, 688.) From feces of in- 
fants. 

Micrococcus communis lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900, 48.) From milk. Hucker (loc. 
cit., 19) considers this a synonym of 
Micrococcus freudenreichii Guillebeauor 
of Micrococcus ureae Cohn. 

Micrococcus commensalis (Turro) Mi- 
gula. (Diplococcus commensalis Turro, 
Cent. f. Bakt., 16, 1894, 1; Migula, Syst. 



d. Bakt., 2, 1900, 125.) From sputum. 
Winslow and Winslow {loc. cit., 220) 
consider this a synonym of Micrococcus 
luteus Cohn. 

Micrococcus commutatus De Toni and 
Trevisan. {Micrococcus albus I or Mi- 
crococcus albus fluidificans Maggiora, 
Giorn. Soc. Ital. d'Igiene, 11, 1889, 350; 
De Toni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1079.) 

Micrococcus concentricus Zimmermann. 
(Die Bakt. unserer Trink- u. Nutz- 
wjisser, Chemnitz, I Reihe, 1890, 86.) 
From water. Winslow and Winslow 
(loc. cit., 224) state that this is appar- 
ently a synonym of Micrococcus candicans 
Fltigge. 

Micrococcus confluens Kern. (Arb. 
bakt. Inst. Karlsruhe, /, Heft 4, 1897, 
494.) From the stomach and intestine 
of the starling {Sturnus vulgaris) and 
the finch {Fringella carduelis). Win- 
slow and Winslow {loc. cit., 216) consider 
this a synonym of Micrococcus flavus 
Trevisan. 

Micrococcus conjunctivae Migula. 
{Micrococcus liquefaciens conjunctivae 
Gombert, Recherches experimentales sur 
les microbes des conjunctives, Mont- 
pellier and Paris, 1889; Migula, Syst. d. 
Bakt., 2, 1900, 115.) From normal hu- 
man conjunctiva. Hucker {loc. cit., 15) 
considers this a synonym of Micrococcus 
albus Schroeter. 

Micrococcus conjunctividis Migula. 
{Micrococcus flavus conjunctivae Gom- 
bert, Recherches experimentales sur les 
microbes des conjunctives, Montpellier 
and Paris, 1889; Migula, Syst. d. Bakt., 
2, 1900, 141.) From normal human con- 
junctiva. Winslow and Winslow {loc. 
cit., 216) consider this a synonym of 
Micrococcus flavus Trevisan; while 
Hucker {loc. cit., 11) regards it as a 
synonym of Micrococcus citreus Migula. 
Micrococcus conoideus Migula. (Staph- 
ylococcus salivarius pyogenes Biondi, 
Ztschr. f. Hyg., 2, 1887, 227 ; Staphylococ- 
cus sialopyus Trevisan, I generi e le 
specie delle Batteriacee, Milan, 1889, 



258 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



32; Micrococcus salivarius -pyogenes 
Freire, Memoire sur la bacteriologie, 
pathog^nie, traitement et prophylaxie 
de la fievre jaune. Rio de Janiero, 1898; 
Abst. in Cent. f. Bakt., I Abt., 26, 1899, 
741; Staphylococcus pyogenes salivarius, 
quoted from Goadby, Trans, of Odon- 
tolog. Society, June, 1899, see Abst. in 
Cent. f. Bakt., I Abt., Ref., 31, 1902, 493; 
Migula, Syst. d. Bakt., 2, 1900, 102.) 
From saliva. Hucker {loc. cit., 12 and 
15) regards this as a synonym of Micro- 
coccus aureus Zopf or of Micrococcus 
albus Schroeter. 

Micrococcus corallinus Cantani. (Can- 
tani. Cent, f . Bakt., I Abt., 23, 1898, 311 ; 
Rhodococcus coralinus (sic) Levine and 
Soppeland, Iowa State Coll. Engineer- 
ing Exp. Sta. Bui. 77, 1926, 22.) From 
dust. Hucker {loc. cit., 25) considers this 
a synonym of Micrococcus roseus Fliigge. 
Levine and Soppeland (loc. cit.) regard 
this as a synonym of Rhodococcus fulvus 
Winslow and Rogers. For a description 
of this species, see Bergey et al.. Manual, 
5th ed., 1939, 253. 

Micrococcus coralloides Zimmermann. 
(Die Bakt. unserer Trink- u. Nutzwasser, 
Chemnitz, II Reihe, 1894, 72.) From 
water. Winslow and Winslow (loc. cit., 
199) state that this species appears to be 
a synonym of Micrococcus albus Zopf; 
while Hucker (loc. cit., 17) considers it a 
synonym of Micrococcus caseolyticus 
Evans. 

Micrococcus corrugatus Migula. 
(Merismopedia mesentericus corrugatus 
Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 
355; Migula, Syst. d. Bakt., 2, 1900, 161.) 
From dust. Winslow and Winslow (loc. 
cit., 216) consider this a synonym of 
Micrococcus flavus Trevisan. 

Micrococcus coryzae (Hajek) Migula. 
(Diplococcus coryzae Hajek, Berliner 
klin. Wochnschr., No. 33, 1888; Migula, 
Syst. d. Bakt., 2, 1900, 63.) From secre- 
tions in acute catarrh. Winslow and 
Winslow (loc. cit., 205) consider this a 
synonym of Micrococcus candidus Cohn 
or of Gaffkya tetragena Trevisan. 



Micrococcus cremoides Zimmermann. 
(Die Bakt. unserer Trink- u. Nutzwasser, 
Chemnitz, I Reihe, 1890, 74.) From 
water. Winslow and Winslow (loc. cit., 
216) consider this a synonym of Micro- 
coccus flavus Trevisan; while Hucker 
(loc. cit., 10) considers it a synonym of 
Micrococcus conglomeratus Migula. 

Micrococcus cremoides albus Dyar. 
(Ann. N. Y. Acad. Sci., 8, 1895, 350). 
From dust. Regarded by Dyar as a 
white form of Micrococcus cremoides 
Zimmermann. 

Micrococcus cremorisviscosi (Hammer 
and Cordes) Bergey et al. (Staphylococcus 
cremoris-viscosi Hammer and Cordes, 
Jour. Dairy Sci., 3, 1920, 291; Bergey et 
al.. Manual, 3rd ed., 1930, 86.) From 
ropy milk. For a description of this 
species, see Bergey et al.. Manual, 5th 
ed., 1939, 254. 

Micrococcus crepuscidum (Ehrenberg) 
Cohn. (Monas crepuscidum Ehrenberg, 
Abhandl. d. Berliner Akad., 1830, 74 and 
1832, 57 ; Cohn, Beitr. z. Biol. d. Pflanzen, 
1 , Heft 2, 1872, 160.) De Toni and Trevi- 
san (in Saccardo, Sylloge Fungorum, 8, 
1889, 1082) list the following as synonyms 
of this species; Protococcus nebidosus 
Kiitzing, Linneae, 8, 1833, 365; Crypto- 
coccus nebulosus Kiitzing, Phycol. gener., 
1845, 147; Cryptococcus natans Kiitzing, 
Spec. Alg., 1849, 146. 

Micrococcus cretaceus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
65.) From cheese. Winslow and Wins- 
low (loc. cit., 224) state that this is appar- 
ently a synonym of Micrococcus candicans 
Fliigge. 

Micrococcus cristatus Glage. (Ztschr. 
f. Fleisch- u. Milch-hygiene, 10, 1900, 
145.) From the surface of wurst and 
similar meat products. 

Micrococcus cruciformis Freire. 
(Compt. rend. Acad. Sci., Paris, 128, 
1899, 1047.) From the stamens and 
pistils of Hibiscus. 

Micrococcus cumidatus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
497; not Micrococcus cuinulatus Chester, 



FAMILY MICROCOCCACEAE 



259 



see Micrococcus tenuissimus INIigula.) 
From the stomach and intestine of the 
yellow-hammer {Emberiza citrinella) and 
of the finch {Fringella carduelis). 
Hucker {loc. cit., 25) regards this as a 
synonym of Micrococcus roseus Fliigge. 

Micrococcus cupularis Migula. (No. 
29, Lembke, Arch. f. Hyg., 29, 1897, 331 ; 
Migula, Syst. d. Bakt., 2, 1900, 211.)From 
feces. Winslow and Winslow {loc. cit., 
216) consider this a synonym of Micrococ- 
cus flavus Trevisan. 

Micrococcus cupuliformis Migula. (No. 
19, Lembke, Arch. f. Hyg., 29, 1897, 325; 
Migula, Syst. d. Bakt., 2, 1900, 213.) 
From feces. Winslow and Winslow {loc. 
cit., 220) consider this a synonym of 
Micrococcus luteus Cohn. 

Micrococcus curtissi Chorine. (Chor- 
ine, Internat. Corn Borer Invest. Chi- 
cago, 2, 1929, 48.) From diseased larvae 
of the corn borer (Pyrausta nubilalis). 
Also virulent to larvae of the flour moth 
{Ephestia kuhniella) and of the bee moth 
(Galleria mellonella). 

Micrococcus cutis communis Sabouraud. 
(Sabouraud, Ann. d. derma tol. et syphil., 
1896, Heft 3; Abst. in Cent. f. Bakt., I 
Abt., 20, 1896, 249; Staphylococcus cutis 
communis Sabouraud, Practique Derma- 
tologique, /, 1903, 714.) From human 
skin especially in alopecia areata, certain 
types of eczema and acne. May be the 
same as Micrococcus epidermidis Hucker. 

Micrococcus cyaneus (Schroeter) Cohn. 
(Bacteridium cyaneum Schroeter, Beitr. 
z. Biol. d. Pflanzen, /, Heft 2, 1872, 122 
and 126; Cohn, ibid., 156; Nigrococcus 
cyaneus Castellani and Chalmers, Man. 
Trop. Med., 3rd ed., 1919, 932.) From 
dust and water. 

Micrococcus cyanogenus Pammel and 
Combs. (Proc. Iowa Acad. Sci., 3, 1895, 
136; see Abst. in Cent. f. Bakt., II Abt., 
2, 1896, 764.) From milk. 

Micrococcus cyclops Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
69.) From Swiss cheese. Winslow and 
Winslow (loc. cit., 224) state that this is 



appai'ently a synonym of Micrococcus 
candicans Fliigge. 

Micrococcus cystiopoeus Miiller-Thur- 
gau. (Cent. f. Bakt., II Abt., 20, 1908, 
464.) From wine. 

Micrococcus cytophagus Merker. 
(Cent. f. Bakt., II Abt., 31, 1912, 589.) 
Found on the leaves of Elodea. Utilizes 
cellulose. Stanier (Bact. Rev., 6, 1942, 
150) thinks that these micrococci were 
microcysts of Sporocytophaga spp. 

Micrococcus dantecH Chester. (Coc- 
cus du rouge de morue, Le Dantec, Ann. 
Past. Inst., 5, 1891, 662; Chester, Man. 
Determ. Bact., 1901, 106.) From red 
salted codfish. Hucker (loc. cit., 25) 
considers this a synonym of Micrococcus 
roseus Fliigge. 

Micrococcus decalvens (Thin) Schroe- 
ter. (Bacterium decalvens Thin, Monats. 
f. prakt. Dermatol., No. 28, 1885; Schroe- 
ter in Cohn, Kryptog. -Flora v. Schlesien, 
3, 1, 1886, 149.) From hair follicles in 
alopecia areata. 

Micrococcus decipiens Trevisan. (Bac- 
terie de Pair, Cornil and Babes, Les 
Bacteries, 1885, 124; Trevisan, I generi e 
le specie delle Batteriacee, Milan, 1889, 
34.) From dust. 

Micrococcus (Streptococcus ?) decolor 
Migula. (No. 22, Lembke, Arch. f. 
Hyg., 26, 1896, 314; Migula, Syst. d. 
Bakt., 2, 1900, 203.) From feces. 
Hucker (loc. cit., 17) considers this a 
synonym of Micrococcus caseolyticus 
Evans. 

Micrococcus deformans Crowe. (Brit. 
Med. Jour., Nov. 27, 1920, 815; Abst. 
in Cent. f. Bakt., I Abt., Ref., 73, 1922, 
84.) From cases of arthritis. A form 
of Micrococcus pyogenes albus according 
to Lehmann and Neumann (Bakt. Diag., 
7 Aufl., 2, 1927, 293). 

Micrococcus delacourianus Roze. 
(Compt. rend. Acad. Sci., Paris, 123, 
1896, 613 and 1323.) From dry rotting 
potatoes. 

Micrococcus dendroporthos Ludwig. 
(Cent. f. Bakt., 10, 1891, 10.) From the 
bark of poplar trees (Poprdus sp.). 



260 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Micrococcus denitrificans Beijerinck. 
(Cent. f. Bakt., II Abt., 25, 1910, 53.) 
From Rochelle salts (sodium potassium 
tartrate). 

Micrococcus dermatogenes Fuhrmann. 
(Cent. f. Bakt., II Abt., 17, 1906, 618.) 
From bottled beer. 

Micrococcus diffluens Schroeter. (In 
Cohn, Kryptog. -Flora v. Schlesien, 3, 1, 
1886, 144.) From dust, feces, etc. 

Micrococcus dimorpkus Bucherer. 
(Planta, Arch. f. wissen. Bot., 1934, 98.) 
A dimorphic bacterium. He reports it 
as much like Micrococcus melitensis 
Bruce and Bacterium fraenkelii Hashi- 
moto. 

Micrococcus diphtericus (sic) Cohn. 
(Micrococcus, Oertel, Deutsch. Arch. f. 
klin. Med., 8, 1871 ; Cohn, Beitr. z. Biol, 
d. Pflanzen, i, Heft 2, 1872, 162; Strepto- 
coccus diphtheriticus Zopf, Die Spalt- 
pilze, 3 Aufl., 1885, 53.) From throats 
and nasal passages of diphtheria patients. 

Micrococcus dissimilis Dyar. (Sec 

Sattler, Cent. f. Bakt., 5, 1889, 70; Dyar, 
Ann. N. Y. Acad. Sci., 8, 1895, 353 ; Micro- 
coccus trachomatis conjunctivae Sattler in 
Krai, Die gegenwartigen Bestand der 
Kral'schen Sammlung von Mikroorgan- 
ismen, 1900, 19.) From trachoma infec- 
tions. Hucker (loc. cit., 17) considers 
this a synonym oi Micrococcus caseolyticus 
Evans. 

Micrococcus djokjakartensis Zettnow. 
(Cent. f. Bakt., I Abt., Orig., 75, 1915, 
376.) From a sugar factory in Java. 

Micrococcus doyeni De Toni and Trevi- 
san. {Micrococcus urinae alhus olearius 
Doyen, Jour. d. connaiss. medic, No. 14, 
1889, 108; De Toni and Trevisan, in 
Saccardo, Sylloge Fungorum, 8, 1889, 
1076.) From urine. Hucker (loc. cit., 
16) states that this species is apparently 
identical with Micrococcus albus 
Schroeter. 

Micrococcus drimophylus Baumgartner. 
(Baumgartner, Ergebnisse d. ges. Zahn- 
heilk.. Heft 2, 1910, 729; Abst. in Cent, 
f. Bakt., I Abt., Ref., 48, 1911, 622.) 
From the mouth cavitv. 



Micrococcus eatonii Corbet. (Quart. 
Jour. Rubber Research Inst. Malaya, 
2, 1930, 145.) From the latex of the 
rubber tree (Hevea hrasiliensis). For 
a description of this species, see Bergey 
et al.. Manual, 5th ed., 1939, 244. 

Micrococcus eburneus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
85.) From Camembert cheese. Win- 
slow and Winslow {loc. cit., 224) state 
that this species is apparently a synonym 
of Micrococcus candicans Flixgge. 

Micrococcus ephestiae Mattes. (Sit- 
zungsber. d. Gesellsch. z. Beford. d. 
gesamt. Naturwissensch. zu Marburg, 
62, 1927, 406.) From the Mediterranean 
flour moth {Ephestia kuehniella). 

Micrococcus epimetheus Corbet {loc. 
cit., 148). From the latex of the rubber 
tree {Hevea hrasiliensis). For a de- 
scription of this species, see Bergey et al.. 
Manual, 5tli ed., 1939, 256. 

Micrococcus esterificans Beck. (Arb. 
kaiserl. Gesundheitsamte, 29, Heft 2, 
1905; Abst. in Cent. f. Bakt., II Abt., 
19, 1907, 594.) Has a characteristic 
fruity aroma. From butter. 

Micrococcus exanthematicus Lewa- 
scheff. (Deutsch. med. Wochnschr., No. 
13 and 34, 1892; Abst. in Cent. f. Bakt., 
12, 1892, 635.1 From blood in cases of 
typhus fever. Motile. Grows anaer- 
obically. 

Micrococcus excavatus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
486.) From the stomach contents of a 
coot {Fulica atra) and a woodpecker 
(Picus major). Winslow and Winslow 
{loc. cit., 220) consider this a synonym of 
Micrococcus luteus Cohn. 

Micrococcus exiguus Kern {loc. cit., 
470). From the stomach contents of the 
chaffinch {Fringella coelebs). Winslow 
and Winslow {loc. cit., 199) state that 
this appears to be a synonym of Micro- 
coccus alMis Schroeter; while Hucker 
(loc. cit., 19) considers it a synonym of 
Micrococcus Jreudenreichii Guillebeau 
or of Micrococcus ureae Cohn. 

Micrococcus expositionis Chester. (No. 



FAMILY MICROCOCCACEAE 



261 



34, Conn, Storrs Agr. Exp. Sta. 7th Ann. 
Rept., 1895, 77; Chester, Man. Determ. 
Bact., 1901, 92.) From air. Winslow 
and Winslow {loc. cit. 216) consider this 
a synonym of Micrococcus flavus Trevi- 
san; while Hucker (loc. cit., 10) regards 
it as a synonym of Micrococcus conglom- 
eratus Migula. 

Micrococcus expressus Weiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
195.) From a carrot infusion. Produces 
slime. Hucker {loc. cit., 7) considers 
this species a synonym of Micrococcus 
luteus Cohn or of Micrococcus varians 
Migula. 

Micrococcus faviformis Migula. 
(Milchweisser Diplococcus, Bumm, Mi- 
kroorg. d. gonorrh. Schleimbautkr., II 
Ausg., 1887, 18; Micrococcus lacteus 
faviformis Fliigge, Die Mikroorganismen, 
2 Aufl., 1886, 182; Neisseria lactea Trevi- 
san, I generi e le specie delle Batteriacee, 
Milan, 1889, 32; Migula, Syst. d. Bakt., 
2, 1900, 117.) From vaginal and other 
body secretions. Winslow and Winslow 
(loc. cit., 199) state that this appears 
to be a synonym of Micrococcus albus 
Schroeter. 

Micrococcus feddei Herter. (Micro- 
coccus XVI, Choukevitch, Ann. Inst. 
Past., 25, 1911, 354; Botan. Jahresber., 
39, II Abt., Heft 4, 1914, 755; Abst. in 
Cent. f. Bakt., II Abt., 51, 1920, 367.) 
From the large intestine of a horse. Re- 
sembles Micrococcus roscidur Matzu- 
schita. 

Micrococcus fervitosus Adametz and 
Wichmann (Bakt. d. Trink- u. Xutz- 
wasser. Mitt. Oest. Versuchstat. f. 
Brauerei u. Malzerei, Wien, Heft 1, 
1888.) From water. Winslow and 
Winslow {loc. cit., 205) consider this a 
synonym of Gaffkya tetragcna Trevisan. 

Micrococcus fickii Trevisan. {Coccus 
albus non liquefaciens {Coccus candicans) 
Fick, Ueber Microorg. in Conjunctival- 
sack, Wiesbaden, 1887; Trevisan, I generi 
e le specie delle Batteriacee, Milan, 1889, 
33.) From conjunctiva. 

Micrococcus finlayensis Sternberg. 
(Rept. on Etiology and Prevention of 



Yellow Fever, Washington, 1891, 219.) 
Obtained by Finlay in cultures from the 
liver and spleen of a yellow-fever cad- 
aver. Hucker {loc. cit., 11) considers 
this a synonym of Micrococcus citreus 
Migula. 

Micrococcus flaccidifex danai Brown. 
(Amer. Museum Novit., No. 251, 1927, 
5.) Causative agent of wilt disease of 
monarch butterfly larvae {Danais archip- 
pus). Considered a sub-species of Gyro- 
coccus flaccidifex Glaser and Chapman 
(Science, 36, 1912, 219). 

Micrococcus flagcllatus Klotz. (Jour. 
Med. Research, 11 (N.S.6), 1904, 493.) 
Found in an epizootic among rabbits and 
white rats. Supposedly flagellated. 

Micrococcus flavens Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
80.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 216) consider this a 
synonym of Micrococcus flavus Trevisan. 

Micrococcus flavescens Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
79.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 216) consider this a 
synonym of Micrococcus flavxis Trevisan. 
For a description of this species, see 
Bergey et al., Manual, 5th ed., 1939, 246. 

Micrococcus flavidus Henrici. {loc 
cit., 81; not Micrococcus flavidus Roze, 
Compt. rend. Acad. Sci., Paris, 122, 1896, 
750.) From Swiss and Limburger 
cheeses. Winslow and Winslow {loc. 
cit., 216) consider this a synonym of 
Micrococcus flavus Trevisan. 

Micrococcus flavovirens Migula. 
{Staphylococcus viridis flavescens Gutt- 
mann, Arch. f. path. Anat., 107, 1887, 
261 ; Staphylococcus viridi-flavescens 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 33; Migula, Syst. 
d. Bakt., 2, 1900, 124; Micrococcus viridis 
Chester, Man. Determ. Bact., 1901, 95; 
Micrococcus viridis -flavescens Winslow 
and Winslow, Systematic Relationships 
of the Coccaceae, 1906, 221.) Winslow 
and Winslow {ibid., 220) consider this a 
synonym of Micrococcus luteus Cohn. 

Micrococcus flavus non liquefaciens 



262 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Amsler. (Amsler, Korrespondenbl. f. 
Schweizer Aerzte, 1900, No. 9; Abst. in 
Cent. f. Bakt., I Abt., 29, 1901, 450.) 
From thermal springs. 

Micrococcus fluorescens Maggiora. 
(Giorn. Soc. Ital. d'Igiene, 11, 1889, 352; 
Abst. in Cent. f. Bakt., 8, 1890, 13.) 
From the skin of the foot. 

Micrococcus foetidus Fliigge. (Die Mi- 
kroorganismen, 2 Aufl., 1886, 172; not 
Micrococcus foetidus Veillon, Compt. 
rend. Soc. Biol. Paris, 1893, 867; see 
Streptococcus foetidus Prevot.) From 
carious teeth. 

Micrococcus foetidus Klamann. (All- 
gem, med. Centralzeitung, 1887, 1344.) 
Isolated from the posterior nares of man. 
Winslow and Winslow (loc. cit., 199) 
state that this appears to be a synonym 
of Micrococcus albus Schroeter. 

Micrococcus frag His (Dyar) Migula. 
(Merismopedia fragilis Dyar, Ann. N. Y. 
Acad. Sci., 8, 1895, 351; Migula, Syst. d. 
Bakt., 2, 1900, 186.) From dust. Muck- 
er (loc. cit., 25) states that this species 
may be identical with M icrococcus roseus 
Fliigge. 

Micrococcus franklandioruin Trevisan. 
(Micrococcus candicans Frankland and 
Frankland, Phil. Trans. Roy. Soc. Lon- 
don, 178, B, 1888, 270; not Micrococcus 
candicans Fliigge, Die Mikroorganismen, 
2 Aufl., 1886, 173; Trevisan, I generi o 
le specie delle Batteriacee, Milan, 1889, 
34.) From dust. 

Micrococcus fulvus Weiss. (Arb. a. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
206; not Micrococcus fulvus Cohn, Beitr. 
z. biol. d. Pflanz., /, Heft 3, 1875, 181.) 
From a bean infusion. 

Micrococcus fuse us Adametz . (Brauner 
Coccus, Maschek, Jahresb. d. Kommunal- 
Oberrealsch. zu Leitmeritz, No. 6, 1867, 
60; Adametz, Bakt. d. Nutz- u. Trink- 
wasser, Vienna, 1S88; Micrococcus fuscus 
Castellani and Chalmers, Man. Trop. 
Med., 3rd ed., 1919, 932.) Hucker (loc. 
cit., 10) states that this species is prob- 
ably identical with Micrococcus con- 
glomeratus Migula. 



Micrococcus galbanatus Zimmermann. 
(Bakt. unserer Trink- u. Nutzwasser, 
Chemnitz, II Reihe, 1894, 68.) From 
water. Winslow and Winslow (loc. cit., 
216) consider this a synonym of Micro- 
coccus flavus Trevisan. 

Micrococcus gallicidus Burrill. (Amer. 
Nat., 17, 1883, 320.) From blood of fowls 
infected with chicken cholera. 

Micrococcus gelatinogenus Briiutigam. 
(Pharmaceutische Centralhalle, 32, 1891, 
427.) From digitalis infusions. See 
Micrococcus gummosus Happ. 

Micrococcus gelatinosus Warrington. 
(The Lancet, 1, 1888, No. 25; Abst. in 
Cent. f. Bakt., 4, 1888, 394.) Curdles 
milk. 

Micrococcus gelatinosus Issatchenko. 
(Recherches sur les Microbes de I'Ocean 
Glacial Arctique, Petrograd, 1914, 232; 
not Micrococcus gelatinosus Warrington, 
The Lancet, 1, 1888, No. 25.) From sea 
water. 

Micrococcus giganteus lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900, 46.) From milk. 

Micrococcus gigas Frankland and 
Frankland. (Philos. Trans. Roy. Soc, 
London, 178, B, 1888, 268.) From dust. 
Winslow and Winslow (loc. cit., 216) con- 
sider this a synonym of Micrococcus 
flavus Trevisan. 

Micrococcus gilvus Losski. (Inaug. 
Diss., Dorpat, 1893, 60.) Winslow and 
Winslow {loc. cit., 220) consider this a 
synonym of Micrococcus luteus Cohn. 

Micrococcus gingivae Migula. (Mi- 
crococcus gingivae pyogenes Miller, Die 
Mikroorganismen d. Mundhohlc, Leipzig, 
1889, 216; Migula, Syst. d. Bakt., 2, 1900, 
68.) From alveolar pyorrhoea, also from 
the mouth of a healthy man. 

Micrococcus gingreardi Renault. 
(Compt. rend. Acad. Sci., Paris, 120, 
1895,217.) 

Micrococcus glandulosus Weiss. (Arb. 
bakt. Inst. Karlsruhe, ^, Heft 3, 1902, 201 . ) 
From an asparagus infusion. Hucker 



FAMILY MICROCOCCACEAE 



263 



{loc. cit., 19) regards this speciee as 
identical with Micrococcus freiidenreichii 
Guillebeau or with Micrococcus ureae 
Cohn. 

Micrococcus globosus Kern. (Arb. 

bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
469.) From the stomach contents of a 
coot {Fulica atra). Winslow and Wins- 
lov,' {loc. cit., 224) state that this is appar- 
ently a synonym of Micrococcus candicans 
Fliigge. 

Micrococcus granulatus Weiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
197.) From a malt infusion. 

Micrococcus granulosus Kern {loc. cit., 
483). From the stomach contents of the 
yellow-hammer {Emheriza citrinella) and 
the starling {Sturnus vulgaris). Wins- 
low and Winslow {loc. cit., 220) consider 
this a synonym of Micrococcus luteus 
Cohn. 

Micrococcus griseus (Warming) Winter. 
{Bacterium griseum Warming, Viden- 
skabelige Meddelelser fra den naturhist. 
Forening i Kjobenhavn, 1875, 398; 
Winter, in Rabenhorst, Kryptog. -Flora 
V. Deutschl., Oesterr. u. d. Schweiz, 2 
Aufi., 1, 1884,47.) 

Micrococcus grossus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
71.) From Camembert cheese. Wins- 
low and Winslow {loc. cit., 224) state 
that this is apparently a synonym of 
Micrococcus candicans Fliigge. 

Micrococcus gummosus Happ. (Happ, 
Inaug. Diss., Basel, 1893, 31; not Micro- 
coccus gummosus Weiss, Arb. bakt. Inst. 
Karlsruhe, 2, Heft 3, 1902, 189.) From 
snakeroot and digitalis infusions. Pre- 
sumably Leuconostoc mesenieroides Van 
Tieghem. 

Micrococcus haematodes Zopf. (Mi- 
crobes de la sueur rouge. Babes, Biol. 
Centralbl., 2, 1882, No. 8; Zopf, Die 
Spaltpilze, 3 Aufl., 1885, 60.) The cause 
of red perspiration. Hucker {loc. cit., 
25) states that this may be a synonym of 
Micrococcus roseus Fliigge. 

Micrococcus haemorrhagicus (Klein) 
Migula. {Staphylococcus haemorrhagicus 



Klein, Cent. f. Bakt., I Abt., 22, 1897, 
81; Migula, Syst. d. Bakt., 2, 1900, 88.) 
Associated with an erythema of the skin 
resembling anthrax. Winslow and Wins- 
low {loc. cit., 199) state that this appears 
to be a synonym of Micrococcus albtis 
Schroeter. 

Micrococcus halensis Lehmann and 
Neumann. {Micrococcus acidi para- 
lactici liquefaciens halensis Kozai, Ztschr. 
f. Hyg., 31, 1899, 374; Lehmann and 
Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 
210; Micrococcus acidi paralactici lique- 
faciens Thiele, Ztschr. f. Hyg., ^6, 1904, 
394.) From milk. Hucker {loc. cit., 17) 
considers this a synonym of Micrococcus 
caseolyticus Evans. 

Micrococcus halophilus Bergey et al. 
(Culture No. 19, Baranik-Pikowsky, 
Cent. f. Bakt., II Abt., 70, 1927, 373; 
Bergey et al.. Manual, 3rd ed., 1930, 89.) 
From sea water. For a description of 
this species, see Bergey et al., Manual, 
5th ed., 1939, 258. 

Micrococcus hauseri (Rosenthal) Mi- 
gula. {Diplococcus hauseri Rosenthal, 
Inaug. Diss., Berlin, 1893, 26; Migula, 
loc. cit., 80.) From the oral cavity. 
Winslow and Winslow {loc. cit., 224) state 
that this species is apparently identical 
with Micrococcus candicans Fliigge. 

Micrococcus helvolus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
77.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 220) consider this to 
be identical with Micrococcus luteus 
Cohn . 

Micrococcus hibiscus Nakahama. (Jour. 
Agr. Chem. Soc. Japan, 16, 1940, 345, 
Eng. Abs., Bull. Agr. Chem. Soc, 16, 
1940, 51.) Isolated from retting kenaf 
{Hibiscus). 

Micrococcus humidus Migula. (Micro- 
coccus No. 2, Adametz, Landwirtsch. 
.lahrb., 18, 1889, 239; Migula, Syst. d. 
Bakt., 2, 1900, 50.) From Emmenthal 
cheese. Winslow and Winslow {loc. 
cit., 224) state that this species is appar- 
ently identical with Micrococcus candi- 
cans Fliigge. 



264 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Micrococcus hydrolhermicus Cronquist. 
(Monatsh. f. prakt. Derm., 36, 1903.) 
Optimum temperature 41 °C. 

Micrococcus hymenophagus Renault. 
(Compt. rend. Acad. Sci., Paris, 120, 
1895, 217.) 

Micrococcus imperatoris Roze. (Compt. 
rend. Acad. Sci., Paris, 122, 1896, 
545.) Isolated from potatoes. Probably 
from dust. 

Micrococcus inconspicuus Henrici. 
(Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 
1894,64.) From Swiss cheese. Winslow 
and Winslow {loc. cit., 224) state that 
this species is apparently identical with 
Micrococcus candicans Fliigge. 

Micrococcus indolicus Christiansen. 
(Ac. Path. Micr. Scand., 18, 1934, 42; 
Staphylococcus asacchnrolyticus var. in- 
dolicus Weinberg, Nativelle and Prevot, 
Les Microbes Anaerobies, 1937, 1023.) 
Strict anaerobe. For description see 
Bergey et al., Manual, 5th ed., 1939, 266. 

Micrococcus influenzae Migula. (Mi- 
kroorganismus II, Fischel, Ztschr. f. 
Heilkunde, 12, 1891; Abst. in Cent. f. 
Bakt., 9, 1891, 611; Migula, Syst. d. 
Bakt., 2, 1900, 90.) From the blood of an 
influenza patient. Winslow and Wins- 
low {loc. cit., 199) state that this appears 
to be identical with Micrococcus albus 
Schroeter. 

Micrococcus insectoruiu Burrill. 
(Burrill, Amer. Nat., 17, 1883, 319; Strep- 
tococcus insectoruni De Toni and Trevi- 
san, in Saccardo, Sylloge Fungorum, 8, 
1889, 1061; Bacillus insectoruiu Smith, 
28th Biennial Kept. Kan. State Bd. 
Agri., 1933, 54.) From the cecal organs 
of the chinch bug (Blissus leucopterus). 

Micrococcus intermedius Stark and 
Scheib. (Jour. Dairy Sci., 19, 1936, 
210.) From butter. 

Micrococcus intertriginis ^eyer. (N.Y. 
Med. Jour., 70, 1900, 873; Abst. in 
Cent. f. Bakt., I Abt., 30, 1901, 434.) 
From a case of erythema intertrigo. 

Micrococcus iris Henrici. (Arb. bakt. 
Inst. Karlsruhe, /, Heft 1, 1894, 67.) 
From Limburger cheese. Winslow and 



Winslow (loc. cit., 224) state that this 
species is apparently identical with 
Micrococcus candicans Fliigge. 

Micrococcus irregularis Weiss. (Arb. 
bakt. Inst. Karisruhe, 2, Heft 3, 1902, 
184.) From an infusion of beans and 
carrots. 

Micrococcus jo7igii Chester. (Staph- 
ylococcus pyogenes bovis de Jong, Cent, f . 
Bakt., I Abt., 25, 1899, QA:-, Staphylococcus 
bovis de Jong, ibid., 67; Chester, Man. 
Determ. Bact., 1901, 95.) Associated 
with suppurative processes in cattle. 
Winslow and Winslow {loc. cit., 220) 
consider this a synonym of Micrococcus 
luteus Cohn. 

Micrococcus lactericeus Freund. (In- 
aug. Diss., Erlangen, 1893, 21; Abst. in 
Cent. f. Bakt., 16, 1894, 640.) From the 
human mouth. Hucker {loc. cit., 21) 
regards this as a synonym of Micrococcus 
candidus Cohn or of Micrococcus epider- 
midis Hucker. 

Micrococcus lacteus Henrici. (Arb. 
bakt. Inst. Karisruhe, 1, Heft 1, 1894, 
74.) From cheese. Winslow and Wins- 
low {loc. cit., 199) state that this appears 
to be a synonym of Micrococcus albus 
Schroeter; while Hucker (loc. cit., 19) 
considers it a synonym of Micrococcus 
freudenreichii Guillebeau or of Micro- 
coccus ureae Cohn. 

Micrococcus lactis Chester. (No. 44, 
Conn, Storrs Agr. Exp. Sta., 7th Ann. 
Rept., 1895, 79; Chester, Man. Determ. 
Bact., 1901, 90.) From milk. Winslow 
and Winslow {loc. cit., 224) state that 
this is apparently a synonym of Micro- 
coccus candicans Fliigge. 

Micrococcus lactis II (Hueppe) Scholl. 
(Quoted from Lohnis, Cent. f. Bakt., II 
Abt., 18, 1907, 141.) From milk. 

Micrococcus lactis acidi Krueger. 
(Cent. f. Bakt., 7, 1890, 494.) From 
milk. 

Micrococcus lactis albus Conn, Esten 
and Stocking. (Storrs Agr. Exp. Sta. 
18th Ann. Rept., 1907, 120.) From milk. 
Hucker {loc. cit., 19) considers this a 
synonym of Micrococcus freudenreichii 



FAMILY MICROCOCCACEAE 



265 



Guillebeau or of Micrococcus ureae Cohn. 

Micrococcus lactis amari von Freuden- 
reich. (Bittere Milch Micrococcus, 
Conn, Cent. f. Bakt., 9, 1891, 653; von 
Freudenreich, Cent. f. Bakt., II Abt., 
13, 1904, 407.) From the udder and 
bitter cream. 

Micrococcus lactis aureus Conn, Esten 
and Stocking (loc. cit., 112). From milk, 
butter, cheese, stable dust. Hucker 
{loc. cit., 7 and 12) regards this as a syn- 
onym of Micrococcus luteus Cohn, of 
Micrococcus varians Migula or of Micro- 
coccus aureus Zopf. 

Micrococcus lactis citreus Conn, Esten 
and Stocking (loc. cit., 102). From milk. 
Hucker (loc. cit., 7) considers this species 
identical with Micrococcus luteus Cohn 
or with Micrococcus varians Migula. 

Micrococcus lactis citronus Conn, Esten 
and Stocking {loc. cit., 117). From slime 
on Camembert cheese. Hucker {loc. 
cit., 12) regards this as a synonym of 
Micrococcus aureus Zopf. 

Micrococcus lactis flavus Conn, Esten 
and Stocking {loc. cit., 109). May be 
identical with Micrococcus aurantiacus 
Cohn. From milk. Hucker {loc. cit., 
7 and 12) states that this may be a syn- 
onym of Micrococcus luteus Cohn, of 
Micrococcus varians Migula or of Micro- 
coccus aureus Zopf. 

Micrococcus lactis fluorescens Conn, 
Estenand Stocking {loc. cit., 120). From 
stable dust. Exhibits a green fluores- 
cence. Hucker {loc. cit., 18) states that 
this species is very similar to Micrococcus 
caseolyticus Evans. 

Micrococcus lactis gigas Conn, Esten 
and Stocking {loc. cit., 116). From milk. 
Hucker {loc. cit., 22) states that this 
species is probably identical with Micro- 
coccus candidus Cohn or Micrococcus 
epidermidis Hucker. 

Micrococcxis lactis giganteus Conn, 
Estenand Stocking {loc. cit., 122). From 
milk. 

Micrococcus lactis minutissimus Conn, 
Esten and Stocking {loc. cit., 119). From 
milk. Hucker (loc. cit., 10) considers 



this a synonym of Micrococcus conglom- 
eratus Migula. 

Micrococcus lactis rosaceus Conn, Esten 
and Stocking (loc. cit., 109). From milk. 
Hucker (loc. cit., 26) states that this is 
probably identical with Micrococcus 
roseus Fliigge. 

Micrococcus lactis rugosus Conn, Esten 
and Stocking (loc. cit., 122). From milk. 

Micrococcus lactis varians Conn, Esten 
;ind Stocking (loc. cit., 121). Com- 
monly found in milk. May be identical 
with Micrococcus aureus Zopf. Hucker 
(loc. cit., 15) states that this may be 
identical in part with Micrococcus albus 
Schroeter. 

Micrococcus lardarius Krassilschtschik. 
(Memoires Soc. Zool. de France, 9, 1896, 
513; Compt. rend. Acad. Sci., Paris, 123, 
1896, 428.) From diseased silkworms. 
Differs from Streptococcus bombycis. 
Hucker {loc. cit., 22) states that this is 
probably identical with either Micrococ- 
cus candidus Cohn or Micrococcus 
epidermidis Hucker. 

Micrococcus lembkei Migula. (No. 21, 
Lembke, Arch. f. Hyg., 29, 1897, 327; 
Migula, Syst. d. Bakt., 2, 1900, 212.) 
From feces. Winslow and Winslow (loc. 
cit., 220) consider this a synonym of 
Micrococcus luteus Cohn, while Hucker 
(loc. cit., 11) regards it as probably identi- 
cal with Micrococcus citreus Schroeter. 

Micrococcus lentus Migula. (Xo. 22, 
Lembke, loc. cit., 328; Migula, loc. cit., 
209.) From feces. Winslow and Wins- 
low (loc. cit., 199) state that this appears 
to be a synonym of Micrococcus albus 
Schroeter; while Hucker (loc. cit., 19) 
regards it as probably identical with 
Micrococcus freudenreichii Guillebeau 
or Micrococcus ureae Cohn. 

Micrococcus licheniformis Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
482.) From the intestine of the yellow- 
hammer (Emberiza citrinella) . Winslow 
and Winslow (loc. cit., 220) consider this 
a synonym of Micrococcus luteus Cohn. 

Micrococcus lignithum Renault. 



266 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



(Compt. rend. Acad. Sci., Paris, 126, 
1898, 1828.) A fossil form from lignite. 

Micrococcus lipolyticus Horowitz- 
Whissowa. (Ztsehr. f. Untersiuth. d. 
Lebensmittel, 62, 1931, 602.) A fat 
splitting micrococcus from salted fish. 

Micrococcus lipolyticus Stark and 
Scheib. (Jour. Dairy Sci., 19, 1936, 
210; not Micrococcus lipolyticus Horo- 
witz-Wlassowa, loc. cil.) From butter. 

Micrococcus liquefaciens Migula. (Mi- 
crococcus ureae liquefaciens Fliigge, Die 
Mikroorganismen, 2 Aufl., 1886, 169; 
Streptococcus nelhebius Trevisan, Igeneri 
e le specie delle Batteriacee, Milan, 1889, 
31; Staphylococcus ureae liquefaciens 
Lundstrora, Festsclir. d. path. anat. 
Inst. z. Andenken a.d. 250 jiihrige Beste- 
hen d. finnland. Univ. z. Helsingfors, 
1890; abst. in Cent. f. Bakt., 9, 1891, 
672; Migula, Syst. d. Bakt., 2, 1900, 106; 
Micrococcus aethebius Chester, Man. 
Determ. Bact., 1901, 77; Urococcus lique- 
faciens flueggei Mirjuel and Cambier. 
Traite de Bact., 1902, 627; Micrococcus 
liquefaciens flueggei Miquel in Lafar, 
Ilandb. d. tech. :\Iykol., S, 1904-06, 75; 
Albococcus ureae Kliglcr, Jour. Inf. Dis., 
12, 1913, 442; not Micrococcus liquefaciens 
Bergey et al., Manual, 1st cd., 1923, 67.) 
From urine. Huckcr (loc. cil., 18 and 
19) states that this species may be identi- 
cal with Micrococcus caseolyticus Evans, 
Micrococcus freudenreichii Guillebeau oi- 
Micrococcus ureae Cohn. 

Micrococcus liquefaciens acidi /and //, 
Conn. (Storrs Agr. Exp. Sta. 12th Ann. 
Kept., 1900, 48.) From milk. Hucker 
(loc. cil., 18) regards this species as very 
similar to Micrococcus caseolyticus Evans. 

Micrococcus liquidus Migula. (No. 21, 
Lembke, Arch. f. Hyg., 26, 1896, 313; 
Migula, Syst. d. Bakt., 2, 1900, 208.) 
From feces. Hucker (loc. cit., 15) states 
that this species appears identical with 
Micrococcus albus Sehroeter. 

Micrococcus lobatus Migula. (Siebert, 
Inaug. Diss., Wiirzburg, 1894, No. 3, 10; 
Migula, Syst. d. Bakt., 2, 1900, 139.) 
From the human scalp. Winslow and 



Winslow (loc. cit., 184) state that this is 
apparently a synonym of Micrococcus 
aureus Zopf. 

Micrococcus loewenbergii Trevisan. 
(Micrococcus de I'ozene, Lowenberg, 
Congres des otologistes, 1884 and Union 
medicale, 1884; Trevisan, I generi e le 
specie delle Batteriacee, Milan, 1889, 
33.) From secretions in ozena. 

Micrococcus luridus Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft, 4, 1897, 480.) 
From the intestine of the chaffinch (Frin- 
gilla coelebs). Winslow and Winslow 
(loc. cit., 220) consider this a synonym 
of Micrococcus luieus Cohn. 

Micrococcus luteolus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
82; not .Micrococcus luteolus Irwin and 
Harrison, Le Lait, 8, 1928, 881.) From 
cheese. Winslow and Winslow (loc. cit., 
216) consider this a synonym of Micro- 
coccus flavus Trevisan. For a description 
of Irwin and Harrison's organism, see 
Bergey et al., Manual, 5th ed., 1939, 
249.) 

Micrococcus luteus var. larvae Haudu- 
ro3" et a\ . (M icrococcus luteus -liquefaciens 
var. larvae Toumanoff, Bull. Soc. Centr. 
de Med. Veter., 80, 1927, 367; Hauduroy 
et al.. Diet. d. Bact. Path., 1937, 277.) 
From foulbrood of bees. Pathogenic. 

Micrococcus lutosus Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 489.) 
From the stomach contents of the chaf- 
finch (Fringilla coelebs). Winslow and 
Winslow (loc. cit., 216) consider this a 
synonym of Micrococcus flavus Trevisan. 

Micrococcus lysodeikticus Fleming. 
(Proc. Roy. Soc. London, Ser. B, 93, 1922, 
306.) Non-pathogenic. 

Micrococcus lyssae (Rivolta) Trevisan. 
(Cocco-baclerium lyssae Rivolta, 1886; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 33.) Spore-bearer. 

Micrococcus madidus Migula. (No. 
19, Lembke, Arch, f . Hyg., 26, 1896, 311 ; 
Migula, Syst. d. Bakt., 2, 1900, 207.) 
From feces. Winslow and Winslow (loc. 
cit., 184) state that this is apparently a 
synonym of M icrococcus aureus Aligula ; 



FAMILY MICROCOCCACEAE 



26'; 



while Hucker {loc. cit., 15) regards it as 
identical with Micrococcus aJbris 
Schroetcr. 

Micrococcus inagnus (Miller) Trevisan. 
{Jodococcus magnus Miller, Deutsche 
mod. Wochnschr., Xo. 30, 1888 ; Trevisan, 
I generi c le specie dellc Batteriacee, 
Milan, 1889, 33; not Micrococcus magnus 
Chester, Man. Determ. Bact., 1901, 85 
( Diplococcus magnus Rosenthal, Cent. f. 
Bakt., 25, 1899, 1); not Micrococcus 
magnus Stark and Scheib, .lour. Dairy 
Sci., 19, 1936, 210.) 

Micrococcus major Dc Toni and Trevi- 
san. (Micrococcus urinae major Doyen, 
Jour. d. connaiss. medic, Xo. 14, 1889, 
108; De Toni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1076.) From 
urine. 

Micrococcus major Eckstein. (Ztschr. 
f. Forst- u. Jagdwesen. 26. 1894, 18; 
not Micrococcus major DeToni and Tre- 
visan, in Saccardo, Sylloge Fungorum, 
8, 1889, 1076.) Isolated from the larvae 
of the nun moth (Lymantria monacha) 
and Hypono77ieuta sp. 

Micrococcus manfredii Trevisan. (Mi- 
crococcus der progressiven Lymphome 
im Tierorper, Manfredi, Fortschr. d. 
Med., 1886, 713; Trevisan, I generi e Ic 
specie delle Batteriacee, Milan, 1889, 
33; Streptococcus manfredii DeToni and 
Trevisan, in Saccardo, Sylloge Fungo- 
rum, 8, 1889, 1056; Micrococcus canus 
Migula, Syst. d. Bakt., 2, 1900, 63.) 
From sputum. Winslow and Winslow 
{loc. cit., 206) regard this as a synonym 
of Micrococcus candidus Colin or of 
Gaffkya tetragena Trevisan. 

Micrococcus malolacticus Scifert. 
(Ztschr. f . d. landwirtsch. Versuchswesen 
in Oesterreich, 1903, 567; Abst. in Cent, 
f. Bakt., II Abt., 10, 1903, 664.) From 
wine. Hucker (loc. cit., 7) considers this 
a synonym of Micrococcus luteus Cohn 
or of Micrococcus varians ISIigula. 

Micrococcus mammitis Hutchens. 
(Hutchens in Besson, Pract. Bact. 
Microbiol, and Serum Therapy, Trans, 
of 5th ed., 1913, 615; not Streptococcus 
mammitis hovis Hutchens, ibid., 613.) 



From gangrenous mammitis of milking 
ewes. This is le microcoque de I'ar- 
aignee or de la mammite gangreneuse, 
Xocard, Ann. Inst. Past., 1, 1887, 417. 

Micrococcus marginaius Wilhelmy. 
(Arb. bakt. Inst. Karlsruhe, 3, 1903, 11 .) 
From meat extract. 

Micrococcus marinus Issatchenko. 
(Recherches sur les Microbes de I'Ocean 
Glacial Arcticjue, Pctrograd, 1914, 147.) 
From sea water. 

Micrococcus mastitidis Holland. 
(Jour. Bact.. 5, 1920, 224.) Nomen 
nudum. 

Micrococcus mastohius Trevisan. 
(Microcoque de la mammite gangreneuse 
des brebis laitieres, Xocard, Ann. Inst. 
Past., 1, 1887, 417; Trevisan, I generi e le 
specie delle Batteriacee, Milan, 1889, 33; 
Micrococcus gangr. avium, quoted from 
Freire, Rio de Janeiro, 1898; Abst. in 
Cent. f. Bakt., I Abt., 26, 1899, 841; 
Micrococcus ovis Migula-, Syst. d. Bakt., 
2, 1900, 90; Micrococcus mastilis Chester, 
Man. Determ. Bact., 1901, 76; Micrococ- 
cus mastidis gangraenosae ovis Pfeiler, 
Ztschr. f. Infektionskr., parasit. Krankh. 
u. Hyg. d. Haustiere, 4, 1908, 132.) Said 
to be the cause of gangrenous mastitis 
in sheep. This species appears to be 
Micrococcus pyogenes Migula. 

Micrococcus melanocyclus Merker. 
(Cent. f. Bakt., II Abt., 41, 1911, 589.) 
See Actinomyces melanocyclus Krainsky. 

Micrococcus melanoglossophorus Spe- 
gazzini. (Fung. Arg. Pug., 4, 18--, 316.) 
From the epithelium of the tongue. 

Micrococcus meldensis Roger. (Ber. 
An. de Soc. d'Agric. de Meau.x, 1898.) 

Micrococcus mclleus grandinis Harri- 
son. (Bot. Gazette, 26, 1898, 211.) 

Micrococcus memelensis Leichmann. 
(Coccus, Leichmann, Cent. f. Bakt., 
II Abt., 2, 1896, 780; Leichmann, in 
Koch, Jahresber., 12, 1901, 254; Micro- 
coccus acidi laevolactici Weigmann, in 
Lafar, Handb. d. techn. Mykol., 2, 1905, 
62.) From milk. 

Micrococcus minimus ^yeiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 



268 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



188; not Micrococcus minimus Bergey et 
al., Manual, 1st ed., 1923, 69.) From a 
bean infusion. Hucker (loc. cit., 7) 
considers this a synonym of either Micro- 
coccus luteus Cohn or Micrococcus varians 
Migula. 

Micrococcus minutissimus Issatclienkij. 
(Recherches sur les Microbes de I'Ocean 
Glacial Artique, Petrograd, 1914, 146.) 
From sea water. 

Micrococcus mirificus (Rabenhorst) 
Trevisan. {Palmella miriUca Raben- 
horst, Hedwigia, 1867, 115, and Flor. 
Europ. Algar., 3, 1856, 35; Trevisan, 
Rendic. R. 1st. Lombardo, 12, 1879.) 

Micrococcus mollis (Dyar) Migula. 
(Merismopedia mollis Dyar, Ann. N. Y. 
Acad. Sci., 8, 1895, 352; Migula, Syst. d. 
Bakt., 2, 1900, 161; Aurococcus mollis 
Winslow and Rogers, Science, 21, 1905, 
669; Staphylococcus mollis Holland, Jour. 
Bact., 5, 1920, 225.) From air. A cause 
of boils in the tropics, according to Castel- 
lani and Chalmers (Man. Trop. Med., 
3rd ed., 1919, 931). Hucker {loc. cit., 
12) states that this species is apparently 
identical with Micrococcus aureus Zopf. 

Micrococcus (Diplococcus) morrhuae 
Klebahn. (Mitteil. Inst. Allgm. Botan. 
Hamburg, 4, 1919, 11-69; Abst. in Cent. 
f. Bakt., II Abt., S£, 1921, 123.) Halo- 
philic. Associated with spoilage of salted 
fish. 

Micrococcus mucilagineus Weiss. 
(Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 
1902, 191.) From bean infusions. 
Hucker (loc. cit., 11) states that this is 
probably a synonym of Micrococcus 
citreus Migula. 

Micrococcus mucilaginosus Migula. 
(Micrococcus der schleimigen Milch, 
Ratz, Arch. f. Tierheilkunde, 12, Heft 1 
and 2, 1890; Migula, Syst. d. Bakt., 2, 
1900, 119; not M icrococcus mucilaginosus 
Weiss, Arb. bakt. Inst. Karlsruhe, 2, 
Heft 3, 1902, 205.) From slimy milk. 
Winslow and Winslow {loc. cit., 199) state 
that this appears to be a synonym of 
Micrococcus albus Schroeter; while 
Hucker {loc. cit., 18) considers it a syn- 



onym in part of Micrococcus caseolyticus 
Evans. 

Micrococcus mucofaciens Thoni and 
Thaysen. (Cent. f. Bakt., II Abt., 36, 

1913, 359; not Micrococcus mucofaciens 
Pribram, Klassifikation der Schizomy- 
ceten, 1933, 42.) From milk. Hucker 
{loc. cit., 9) considers this a synonym of 
Micrococcus flavus Trevisan. For a de- 
scription of this species, see Bergey et al.. 
Manual, 5th ed., 1939, 245. 

Micrococcus myceticus Castellani. 
(Proc. Soc. Exp. Biol, and Med., 25, 
1928, 535-536.) From gummy lesions. 

Micrococcus mycodermatus Holland. 
(Jour. Bact., 5, 1920, 224.) 

Micrococcus nacreaceus Migula. (Perl- 
rauttergliinzender Diplococcus, Tataroff, 
Inaug. Diss., Dorpat, 1891, 70; Migula, 
Syst. d. Bakt., 2, 1900, 62.) Winslow 
and Winslow {loc. cit., 224) state that 
this is apparently a synonym of Micro- 
coccus candicans Fliigge. 

Micrococcus neoformans Doyen. (Doy- 
en, Le Micrococcus neoformans et les 
n^oplasmes, Paris, 1903.) From can- 
cerous tissue. Shown by Andrewes and 
Gordon (35th Ann. Rept. Local Govt. 
Board, London, 1905-06, 553) to be iden- 
tical with Micrococcus epidermidis albus 
Welch. 

Micrococcus neurotomae Paillot. 
(Compt. rend. Acad. Sci., Paris, 178, 1924, 
246.) Gram-negative. From the larvae 
of Neurotoma nemoralis. 

Micrococcus neuvillei Trevisan. (Mi- 
crococcus G, Malapert-Neuville, 1887; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889,34.) From mineral 
water. 

Micrococcus nigrescens Castellani. 
(Brit. Jour, of Dermatology, 23, 1911, 
341 ; Nigrococcus nigrescens Castellani 
and Chalmers, Man. Trop. Med., 3rd ed., 
1919, 2103.) Produces a black pigment. 
Found in the black variety of trichomy- 
cosis axillaris, a tropical disease. 

Micrococcus nigrofaciens Northrup. 
(Mich. Agr. Exp. Sta. Tech. Bull. No. 18, 

1914, 12; also in Cent. f. Bakt., II Abt., 



FAMILY MICROCpCCACEAE 



269 



41, 1914, 326.) From diseased larvae 
of the June beetle {Lachnosterna sp.) 
and other insects. ^ 

Micrococcus nitidus Kern. (Arb. bakt . 
Inst. Karlsruhe, 1, Heft 4, 1897, 476.) 
From the stomach and intestine of birds. 
Winslow and Winslow {loc. cit., 199) 
state that this appears to be a synonym 
of Micrococcus albus Schroeter; while 
Hucker {loc. cit., 19) regards it as a syn- 
onym of Micrococcus freudenreichii Guil- 
lebeau or of Micrococcus ureae Cohn. 

Micrococcus nitrificans Bergey et al. 
(Micrococcus 6, Rubentschick, Cent. f. 
Bakt., II Abt., 72, 1927, 125; Bergey et 
al.. Manual, 3rd ed., 1930, 88; not Micro- 
coccus nitrificans van Tieghem, Traite 
de Botanique, Paris, 1883.) From sew- 
age filter beds. For a description of this 
species, see Bergey et al., Manual, 5th 
ed., 1939, 257. 

Micrococcus nivalis Chester. (No. 47, 
Conn, Storrs Agr. Exp. Sta. 7th Ann. 
Rept., 1895, 80; Chester, Man. Determ. 
Bact., 1901, 90.) From dust. Winslow 
and Winslow (loc. cit., 224) state that 
this is apparently a sj^nonym of Micro- 
coccus candicans Fltigge. 

Micrococcus niveus Henrici. (Arb. 

bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
66.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 224) state that this is 
apparently a synonym of Micrococcus 
candicans Fliigge. 

Micrococcus nonfermentans Steinhaus. 
(Jour. Bact., 42, 1941, 779.) From the 
alimentary tract of the lyreman cicada 
{Tibicen linnei) and of an unidentified 
damsel fly {Coenagrionidae) . 

Micrococcus nubilis Migula. (Coccus 
B, Foutin, Bakt. Untersuch. von Hagel, 
Wratsch, No. 49 and 50, 1889; see Cent, 
f. Bakt., 7, 1890, 373; Migula, Syst. d. 
Bakt., 2, 1900, 60 ; M icrococcus beta Ches- 
ter, Man. Determ. Bact., 1901, 87.) Iso- 
lated from hail. Winslow and Winslow 
{loc. cit., 205) consider this to be a syn- 
onym of Micrococcus candidus Cohn or of 
Gaffkya tetragena Trevisan. 

Micrococcus nuclei Roze. (Compt. 



rend. Acad. Sci., Paris, 122, 1896, 544.) 
From potatoes. 

Micrococcus obscoenus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
473.) From the stomach contents of a 
crow {Corvus corone). Winslow and 
Winslow {loc. cit., 199) state that this 
appears to be a synonym of Micrococcus 
albus Schroeter; while Hucker (loc. cit., 
19) considers it a synonym of Micrococcus 
freudenreichii Guillebeau or of Micro- 
coccus ureae Cohn. 

Micrococcus ochraceus Rosenthal. (In- 
aug. Diss., Erlangen, 1893, 22; Abst. in 
Cent. f. Bakt., 16, 1894, 1024; not Micro- 
coccus ochraceus Hansgirg, Oestr. Bot. 
Ztschr., 1885, No. 4.) From the oral 
cavity. Winslow and Winslow {loc. cit., 
220) consider this a synonym of Micro- 
coccus luteus Migula. For a description 
of this species see Bergey et al., Manual, 
5th ed., 1939, 242. 

Micrococcus ochroleiicus Prove. (Prove, 
Beitr. z. Biol. d. Pflanz., 4, Heft 3, 1887, 
409; Streptococcus ochroleucus Trevisan, 
I generi e le specie delle Batteriacee, 
Milan, 1889, 31; Planococcus ochroleucus 
Migula, Syst. d. Bakt., 2, 1900, 272.) 
Fromi urine. Motile. 

Micrococcus odoratus Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
73.) From cheese. Winslow and Wins- 
low {loc. cit., 224) state that this is appar- 
ently a synonym of Micrococcus candicans 
Flugge. 

Micrococcus odorus Henrici {loc. cit., 
72). From cheese. Winslow and Wins- 
low {loc. cit., 224) state that this species 
is apparently a synonym of Micrococcus 
candicans FHigge. 

Micrococcus olearius DeToni and 
Trevisan. {Micrococcus urinae flavus 
oZear jMsDoyen, Jour. d. connaiss. Medic, 
No. 14, 1889, 108; DeToni and Trevisan, 
in Saccardo, Sylloge Fungorum, 8, 1889, 
1077.) From urine. Hucker {loc. cit., 
12) considers this species a synonym of 
Micrococcus aureus Zopf. 

Micrococcus olens Henrici. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 1, 1894, 87.) 



270 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



From Swiss cheese. Winslow and Wins- 
low {loc. cit., 216) consider this a syn- 
onym of Micrococcus flavus Trevisan. 

Micrococcus opalescens DeToni and 
Trevisan. (Micrococcus albus II, Mag- 
giora, Giorn. Soc. Ital. d'Igiene, 11, 1889, 
351; DeToni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1078.) 

Micrococcus orbicularis Chester. {Mi- 
crococcus orbicularis flavus Ravenel, 
Mem. Nat. Acad. Sci.,8, 1896, 8; Chester, 
Man. Determ. Bact., 1901, 101.) From 
soil. Winslow and Winslow {loc. cit., 
216) consider this a synonym of Micro- 
coccus flavus Trevisan; while Hucker 
{loc. cit., 10) regards it as a synonym of 
Micrococcus conglomeratus Migula. 

Micrococcus orbiculatus Wright. 
(Mem. Nat. Acad. Sci., 7, 1895, 432.) 
From Schuylkill River water. Winslow 
and Winslow {loc. cit., 220) consider this 
a synonym of Micrococcus luteus Cohn. 

Micrococcus ovalis Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 500; 
not Micrococcus ovalis Escherich, Die 
Darmbakterien des Sauglings, Stuttgart, 
1886, 90.) From the stomach contents 
of the rock dove {Columba livia). 
Hucker (loc. cit., 9) regards this as a 
synonym of Micrococcus flavtis Trevisan. 

Micrococcus fallens Henrici. (Arb. 
bakt. Inst. Karlsruhe, /, Heft 1, 1894, 
61.) From cheese. Winslow and Wins- 
low {loc. cit., 205) consider this a syn- 
onym of Micrococcus candidus Cohn or of 
Gaffkya tetragena Trevisan. 

Micrococcus -pallidus Henrici {loc. 
cit., 62). From cheese. Hucker {loc. 
cit., 7) regards this species as identical 
with either Micrococcus luteus Cohn or 
Micrococcus varians Migula. 

Micrococcus pannosus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
466.) From the stomach contents of the 
rock dove {Columba livia) and the in- 
testine of another dove {Columba oenas). 
Winslow and Winslow {loc. cit., 224) 
state that this is apparently a synonym 
of Micrococcus candicans Fliigge. 

Micrococcus paraffinae Sohngen. 



(Cent. f. Bakt., II Abt., S7, 1913, 595.) 
From garden earth. 

Micrococcus parotitidis Korentschew- 
sky. (Cent. f. Bakt., I Abt., Orig., U, 
1907, 402.) Isolated from cases of paro- 
titis epidemica. 

Micrococcus parvus (Miller) Trevisan. 
{Jodococcus parvus Miller, Deutsche 
med. Wchnschr., No. 30, 1888; Trevisan, 
I generi e le specie delle Batteriacee, 
Milan, 1889, 33.) 

Micrococcus parvus Migula. (No. 14, 
Lembke, Arch. f. Hyg., 26, 1896, 309; 
Migula, Syst. d. Bakt., 2, 1900, 200.) 
From feces- Winslow and Winslow {loc. 
cit., 224) state that this species is appar- 
ently a synonym of Micrococcus candi- 
cans Fliigge. 

Micrococcus pasteuri Trevisan. (Mi- 
crobe pyogene de I'eau de Seine, Pasteur, 
1877; Trevisan, I generi e le specie delle 
Batteriacee, Milan, 1889, 34; not Micro- 
coccus pasteuri Sternberg, Trans. Pathol. 
Soc. of Philadelphia, 12, 1885, 162.) 
From water. 

Micrococcus pellucidus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
468; not Micrococcus pellucidus Roze, 
Compt. rend. Acad. Sci., Paris, 122, 
1896, 1012.) From the intestine of a spar- 
row {Passer montanus) . Hucker {loc. cit., 
23) regards this as a synonym of Micro- 
coccus candidus Cohn or of Micrococcus 
epidermidis Hucker. 

Micrococcus pemphigiMlgula.. (Diplo- 
coccus des Pemphigus acutus, Demme, 
Verhandl. d. Kongr. f. innere Med., 
Wiesbaden, 1886, 336; Diplococcus pem- 
phigi acuti Lehmann and Neumann, 
Bakt. Diag., 1 Aufl., 2, 1896, 173; Migula, 
Syst. d. Bakt., 2, 1900, V and 191; Micro- 
coccus demmei Chester, Man. Determ. 
Bact., 1901, 74.) Isolated from bullae 
in a case of pemphigus acutus. 

Micrococcus pemphigicontagiosi Castel- 
lani and Chalmers. {Micrococcus pem- 
phigi contagiosa Clegg and Wherry, Jour. 
Inf. Dis., 3, 1906, 171; Castellani and 
Chalmers, Man. Trop. Med., 3rd ed., 
1919, 931 .) From bullae in a case of pem- 



FAMILY MICROCOCCACEAE 



271 



phigus contagiosa. This may be a 
synonym of Micrococcus pemphigi neona- 
torum, see below. 

Micrococcus pemphigineonatorum Cas- 
tellani and Chalmers. {Micrococcus 
pemphigi neonatorum Almquist, Ztsclir. 
f. Hyg., 10, 1891, 253; Staphylococcus 
pemphigi neonatorum Lehmann and Neu- 
mann, Bakt. Diag., 1 Aufl., 2, 1896, 173; 
Castellani and Chalmers, Man. Trop. 
Med., 3rd ed., 1919, 931.) Found in 
bullae in a case of pemphigus neonatorum. 
This may be Micrococcus mollis, accord- 
ing to Castellani and Chalmers {loc. cit.). 
Falls (Jour. Inf. Dis., 20, 1917, 97) identi- 
fies this and the previous organism as 
Micrococcus pyogenes var. aureus Zopf. 

Micrococcus percitreus Bergey et al. 
(Manual, 1st ed., 1923, 63.) From air 
and water. Hucker {loc. cit., 10) con- 
siders this a synonym of Micrococcus 
conglomeratus Migula. For a description 
of this species, see Bergey et al., Manual, 
5th ed., 1939, 248. 

Micrococcus perUavus Bergey et al. 
(Manual, 1st ed., 1923, 62.) From air 
and water. Hucker {loc. cit., 12) re- 
gards this as a synonym of Micrococcus 
aureus Zopf. For a description of this 
species, see Bergey et al.. Manual, 5th 
ed., 1939, 247. 

Micrococcus persicus Kern. (Arb. 

bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
499.) From the intestine of a dove {Co- 
lumba oenas). Hucker {loc. cit., 25) 
states that this may be identical with 
Micrococcus roseus Fliigge. 

Micrococcus petechialis Trevisan. (Mi- 
crococco del dermotifo, Bareggi, 1886; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 33.) 

Micrococcus petilus Trevisan. (Mi- 
crococcus der Pyaemie bei Kaninchen, 
Koch, tjber d. Aetiolog. d. Wundinfec- 
tionskr., Leipzig, 1878 ; Micrococcus pyae- 
miae cuniculorum Schroeter, in Cohn, 
Kryptogam. Flora v. Schlesien, 3, 1, 
1886, 148; Trevisan, I generi e le specie 
delle Batteriacee, Milan, 1889, ZZ; Micro- 



coccus cuniculorum Migula, Syst. d. 
Bakt., 2, 1900, 192.) From rabbits. 

Micrococcus peiroZez Renault. (Compt. 
rend. Acad. Sci., Paris, m, 1897, 1315.) 
A fossil form from oil bearing rocks. 

Micrococcus pieridis Burrill. (Quoted 
from Chittenden, U. S. Dept. Agr., 
Farmers' Bull. No. 1461, 1926, 6.) From 
larvae of the cabbage butterfly {Pieris 
rapae). 

Micrococcus pikowskyi Bergey et al. 
(Culture No. 22, Baranik-Pikowsky, 
Cent. f. Bakt., II Abt., 70, 1927, 373; 
Bergey et al.. Manual, 3rd ed., 1930, 78.) 
From sea water. For a description of 
this species, see Bergey et al., Manual, 
5th ed., 1939,242. 

Micrococcus piliformis Weiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
194.) From a bean infusion. Hucker 
{loc. cit., 7) considers this a synonym of 
Micrococcus luteus Cohn or of Micrococ- 
cus varians Migula. 

Micrococcus piltonensis Gray and 
Thornton. (Cent. f. Bakt., II Abt., 73, 
1928, 81.) From manure and soil. For 
a description of this species, see Bergey 
et al.. Manual, 5th ed., 1939, 259. 

Micrococcus pituitoparus (Hohl) Bu- 
chanan and Hammer. {Karphococcus 
{Carphococcus) pituitoparus Hohl, Jahrb. 
d. Schweiz, 22, 1906, 439; Diplococcus 
viscosus Sato, Cent. f. Bakt., II Abt., 
19, 1907, 27; Buchanan and Hammer, 
Iowa Agr. Exp. Sta. Res. Bull. 22, 1915, 
285.) From slimy milk and from straw. 
Hucker {loc. cit., 23) states that this 
species is probably identical with Micro- 
coccus candidus Cohn or with Micrococcus 
epidermidis Hucker. For a description 
of this species, see Bergey et al., Manual, 
5th ed., 1939, 243. 

Micrococcus plumosus Eisenberg. 
(Brautigam, Inaug. Diss., Leipzig, 1886, 
18; Federiger Micrococcus, Adametz, 
Mitteil. d. Oesterr. Versuchssta. f. Brau- 
erei u. Malzerei, Wien, Heft 1, 1888; 
Eisenberg, Bakt. Diag., 3 Aufl., 1891, 
56.) From feces of cattle and from 
water. Winslow and Winslow {loc. cit., 



272 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



220) consider this a synonym of Micru- 
coccus luteus Cohn; while Hucker {loc. 
cit., 22 and 23) regards it as probably 
identical with Micrococcus candidus Cohn 
or Micrococcus epidermidis Hucker. 

Micrococcus polypus Migula. (Syst. 
d. Bakt., 2, 1900, 79.) From air. 
Hucker {loc. cit., 23) states that this 
species is probably identical with Micro- 
coccus candidus Cohn or Micrococcus 
epidermidis Hucker. 

Micrococcus popxdi Delacroix. (Bui. 
Mens. Off. Renseig. Agr., Paris, 5, 1906, 
1349 and Ann. Inst. Nat. Agron., 2 Ser., 
S, 1906, 353.) Parasitic on poplar trees 
{Popuhis spp.). 

Micrococcus porcellorum Trevisan. 
(Micrococcus bei Hepatitis enzootica 
porcellorum, Nonewitsch, Cent. f. Bakt., 
3, 1888, 233 ; Trevisan, I generi e le specie 
delle Batteriacee, Milan, 1889, 33.) From 
an infected liver. 

Micrococcus progrediens Schroeter. 
(Micrococcus der progressiven Abscess- 
bildung bei Kaninchen, Koch, Uber d. 
Aetiolog. d. Wundinfectionskrankheiten, 
Leipzig, 1878; Schroeter, in Cohn, Kryp- 
togam. -Flora v. Schlesien, 3, 1, 1886, 
148; Micrococcus haernatosaprus Trevi- 
san, I generi e le specie delle Batteriacee, 
Milan, 1889, 33.) From the blood of 
diseased rabbits. 

Micrococcus psalteri Buemann. (Cent. 
f. Bakt., I Abt., Orig., 71, 1913, 308.) 
From the third stomach of cattle. 

Micrococcus pseudocyaneus Schroeter. 
(Kryptogam.- Flora v. Schlesien, 3, 1, 
1886, 145.) A synonym of Micrococcus 
cyaneus Cohn according to Migula, Syst. 
d. Bakt.,^, 1900, 188. 

Micrococcus pseudoinfluenzae Migula. 
(Microorganismus I, Fischel, Ztschr. f. 
Heilkunde, 12, 1891; See Cent. f. Bakt., 
9, 1891, 611; Migula, Syst. d. Bakt., 2, 
1900, 86.) From the blood of an influ- 
enza patient. Hucker (loc. cil., 23) 
considers this a synonym of Micrococcus 
candidus Cohn or of Micrococcus epi- 
dermidis Hucker. 
Micrococcus pulcher Glage. (Ztschr. 



f. Fleisch- u. Milchhyg., 10, 1900, 146; 
not Micrococcus pulcher Weiss, Arb. bakt. 
Inst. Karlsruhe, 2, Heft 3, 1902, 182.) 
From coating on surface of wurst and 
similar meat products. 

Micrococcus pultiformis Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
474.) From stomach contents of the 
yellow-hammer (Emberiza citrinella) and 
starling {Sturnus vulgaris) and from the 
intestine of the woodpecker {Picus ma- 
jor). Winslow and Winslow {loc. cit., 
199) state that this appears to be a syn- 
onym of Micrococcus alb us Schroeter; 
while Hucker {loc. cit., 19) regards it as 
probably identical with Micrococcus 
freudenreichii Guillebeau or with Micro- 
coccus ureae Cohn. 

Micrococcus punctatus Migula. (No. 
18, Lembke, Arch. f. Hyg., 29, 1897, 325; 
Migula, Syst. d. Bakt., 2, 1900, 213.) 
From feces. Winslow and Winslow {loc. 
cit., 199) state that this species appears 
to be a synonym of Micrococcus albus 
Schroeter. 

Micrococcus purpurifaciens Lehmann 
and Neumann. (Micrococcus, Dudt- 
schenko. Cent, f . Bakt., II Abt., 4^, 1915, 
529; Lehmann and Neumann, Bakt. 
Diag., 6 Aufl., 2, 1920, 755.) From ice. 
Produces a purple pigment in alkaline 
gelatin media. 

Micrococcus p^istulatus Henneberg. 
(Cent. f. Bakt., II Abt., 65, 1922, 251.) 
From the human intestine. 

Micrococcus putridus Tilanus. 

(Munch. med.Wchnschr., 34, 1887, 310.) 
From gelatin, agar, etc., containing iodi- 
form. 

Micrococcus pygmaexis Henneberg {loc. 
cit., 252). From the human intestine. 

Micrococcus pyocyaneus Francisco. 
(Revista Valenciana de Ciencias Medicas, 
1914, 2; Abst. in Cent. f. Bakt., I Abt., 
Ref., 63, 1915, 44; not Micrococcus pyo- 
cyaneus Gessard, Thesis, Paris, 1882.) 
From an acne pustule. 

Micrococcus pyosepticus (Hericourt and 
Richet) Solowjew. {Staphylococcus pyo- 
septicus Hericourt and Richet, Compt. 



FAMILY MICROCOCCACEAE 



273 



rend. Acad. Sci., Paris, 107, 1888, 691; 
Solowjew, Abst. in Cent. f. Bakt., I 
Abt., 18, 1895, 60.) From an abscess in 
a dog and from dust. Regarded as identi- 
cal with Micrococcus albus Schroeter. 

Micrococcus quadrigeminus Klebs. 
{Staphylococcus quadrigeminus Vanselow 
and Czaplewski, Cent. f. Bakt., I Abt., 
25, 1899, 143 ; see Lehmann and Neumann, 
Bakt. Diag., 2 Aufl., 2, 1899, 174.) 
Closely related to Micrococcus albus 
Schroeter. 

Micrococcus quaternus Migula. (Sie- 
bert, Inaug. Diss., Wiirzburg, No. I, 
1894, 7; Migula, Syst. d. Bakt., 2, 1900, 
92.) Winslow and Winslow {loc. cit., 
199) state that this appears to be a syn- 
onym of Micrococcus albus Schroeter. 

Micrococcus radiatus Fliigge. (Die 
Mikroorganismen, 2 Aufl., 1886, 176; 
Streptococcus radiatus Crookshank, Man. 
of Bact., 3rd ed., 1890, 256; not Micro- 
coccus radiatus Kern, see below.) From 
dust and water. Winslow and Winslow 
{loc. cit., 199) state that this appears to 
be a synonym of Micrococcus albus 
Schroeter; while Hucker {loc. cit., 18) 
considers it a synonym of Micrococcus 
caseolyticus Evans. 

Micrococcus radiatus Kern. (Kern, 
Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 
1897, 471 ; Micrococcus radiosus Migula, 
Syst. d. Bakt., 2, 1900, 114.) From the 
stomach contents of the starling {Sturnus 
vulgaris). Winslow and Winslow (loc. 
cit., 199) state that this appears to be a 
synonym of Micrococcus albus Schroeter. 

Micrococcus reessii Rosenthal. 
(Inaug. Diss., Berlin, 1893, 19; Abst. in 
Cent. f. Bakt., 16, 1894, 1024.) From 
the oral cavity. Winslow and Winslow 
{loc. cit., 199) state that this appears to 
be a synonym of Micrococcus albus 
Schroeter. 

Micrococcus regular is Weiss. (Arb. 
bakt. Inst. Karlsruhe,^, Heft 3, 1902, 183.) 
From bean infusions. Hucker {loc. cit., 
7) considers this a synonym of Micrococ- 
cus luteus Cohn or Micrococcus varians 
Migula. 



Micrococcus resinaceus Kern. (Arb. 
bakt. Inst. Karlsruhe, /, Heft 4, 1897, 
487.) From the stomach contents of the 
starling {Sturnus vulgaris) and from the 
intestine of a sparrow {Passer montanus). 
Winslow and Winslow {loc. cit., 220) re- 
gard this as a synonym of Micrococcus 
luteus Cohn. 

Micrococcus rhenanus Migula. (Neuer 
Mikrococcus aus Rheinwasser, Burri 
Arch. f. Hyg., 19, 1893, 34; Migula, Syst 
d. Bakt., 2, 1900, 109; Micrococcus rhen 
Chester, ]\Ian. Determ. Bact., 1901, 82 
Albococcus rhejianus Winslow and Rogers, 
Jour. Inf. Dis., 3, 1906, 544.) From 
Rhine River water. Winslow and Wins- 
low {loc. cit., 199) state that this appears 
to be a synonym of Micrococcus albus 
Schroeter; while Hucker {loc. cit., 18) 
considers it a synonym of Micrococcus 
caseolyticus Evans. 

Micrococcus ridleyi Corbet. (Quart. 
Jour. Rubber Research Inst., Malaya, 2, 
1930, 146.) From the latex of the rubber 
tree {Hevea brasiliensis) . For a de- 
scription of this species, see Bergey et 
al., Manual, 5th ed., 1939, 244. 

Micrococcus rosaceus Frankland and 
Frankland. (Trans. Roy. Society, 
London, 178, B. 188, 269; Rhodococcus 
rosaceus Holland, Jour. Bact., 5, 1920, 
225.) From air. Hucker {loc. cit., 25) 
states that this species may be identical 
with Micrococcus roseus Fliigge. For a 
description of this species, see Bergey 
et al.. Manual, 5th ed., 1939, 252. 

Micrococcus rosaceus lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900, 34; Micrococcus lactis rosaceus 
Conn, Esten and Stocking, Storrs Agr. 
Exp. Sta. Rept. for 1906, 108.) From 
milk. 

Micrococcus roscidus Migula. (Micro- 
coccus No. I, Adametz, Landwirtsch. 
.lahrb., 18, 1889, 238; Migula, Syst. d. 
Bakt., 2, 1900, 68.) From Emmenthal 
cheese. Winslow and Winslow {loc. cit., 
224) state that this is apparently a 
synonymofMi'croccccwscandiccns Fliigge. 

Micrococcus roseo-persicinus Migula. 



274 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



(Rote Kokken von Van Ermengem, 
Schneider, Arb. bakt. Inst. Karlsruhe, 
1, Heft 2, 1894, 216; Migula, Syst. d. 
Bakt., 2, 1900, 184.) 

Micrococcus rosettaceus Zimmermann. 
(Bakt. unserer Trink- u. Nutzwasser, 
Chemnitz, I Reihe, 1890, 72.) From 
water. Winslow and Winslow {loc. cit., 
224) state that this is apparently a 
synonym of Micrococcus candicans 
Flugge. 

Micrococcus 7-oseus Maggiora. (Giorn. 
Soc. Ital. d'Igiene, 11, 1889, 356; not 
Micrococcus roseus Fliigge, Die Mikro- 
organismen, 2 Aufl., 1886, 183; not Mi- 
crococcus roseus Gruber, Cent. f. Bakt., 
II Abt., 22, 1909, 408.) 

Micrococcus rubellus Migula. (Syst. d. 
Bakt., 2, 1900, 169.) Source not given. 
Hucker {loc. cit., 27) regards this as 
identical with Micrococcus cinnabareus 
Flugge. 

Micrococcus rubescens Migula. (Xo. 
20, Lembke, Arch. f. Hyg., 26, 1896, 312; 
Migula, Syst. d. Bakt., 2, 1900, 208; 
not Micrococcus rubescens Chester, see 
Micrococcus subroseus below.) Frorh 
feces. Hucker {loc. cit., 27) regards this 
species as identical with Micrococcus 
cinnabareus Flugge. 

Micrococcus rubidus lactis Conn. 
(Conn, Storrs Agr. Exp. Sta. 12th Ann. 
Rept., 1900, 34; Micrococcus lactis rubi- 
dus Conn, Esten and Stocking, Storrs 
Agr. Exp. Sta. 18th Ann. Rept., 1907, 
117.) From milk. Resembles Micro- 
coccus cinnabareus Fliigge. Hucker {loc. 
cit., 25) thinks this species may be iden- 
tical with Micrococcus roseus Fliigge. 

Micrococcus rubigerwsus Kern. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 
492.) From the stomach contents of a 
dove {Columba oenas). Hucker {loc. 
cit., 25) states that this species may be 
identical with Micrococcus roseus Fliigge. 

Micrococcus rubiginosus Passer, and 
Beltr. (Fung. Sicil., 18—, no. 35; quoted 
from DeToni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1082.) 

Micrococcus rugatus Migula. {Micro- 



coccus endocarditidis rugatus Weichsel- 
baum, Beitr. z. path. Anat. u. z. allgm. 
Pathol., 4, 1889, 164; Migula, Syst. d. 
Bakt., 2, 1900, 190; Micrococcus endo- 
carditis Chester, Man. Determ. Bact., 
1901, 74.) From ulcerative endocarditis. 
Winslow and Winslow {loc. cit., 205) con- 
sider this a synonym of Micrococcus can- 
didus Cohn or of Gaffkya tetragena Tre- 
visan; while Hucker {loc. cit., 15) regards 
it as a synonym of Micrococcus albus 
Schroeter. 

Micrococcus rugosus Chester. (No. 2, 
Conn, Storrs Agr. Exp. Sta. 6th Ann. 
Rept., 1894, 50; Chester, Man'. Determ. 
Bact., 1901, 101.) From milk and 
ripened cream. Winslow and Winslow 
{loc. cit., 216) consider this a synonym of 
Micrococcus flavus Trevisan. 

Micrococcus ruminantium Henneberg. 
(Cent. f. Bakt., II Abt., 55, 1922, 252.) 
From the human intestine. 

Micrococcus rushrnorei Brown. (Amer. 
Museum Novit., No. 251, 1927, 4.) Iso- 
lated from a fly {Lucilia sericata) which 
was infected with Bacillus lutzae. 

Micrococcus saccatus Migula. {Micro- 
coccus albus liquefaciens von Besser, 
Beitr. z. i)ath. Anat., 6, 1889, 46; Micro- 
coccus liquefaciens albus, see Cent. f. 
Bakt., 7, 1890, 152; Migula, Syst. d. 
Bakt., 2, 1900, 117; Micrococcus lique- 
faciens Chester, Man. Determ. Bact., 
1901, 78; not Micrococcus liquefaciens 
Holland, Jour. Bact., 5, 1920, 224; Mi- 
crococcus alvi Chester, loc. cit., 81.)From 
the nasal mucous membrane. Winslow 
and Winslow {loc. cit., 199) state that 
this is apparently a synonym of Micro- 
coccus albus Schroeter; while Hucker 
{loc. cit., 19) regards it as probably identi- 
cal with Micrococcus freudenreichii Guille- 
beau or with Micrococcus ureae Cohn. 
For a description of this species, see 
Bergey et al.. Manual, 5th ed., 1939, 254. 

Micrococcus salivalis septicus, quoted 
from Wigura, see Cent. f. Bakt., I Abt., 
17, 1895, 899. From the human skin. 

Micrococcus sarcinoides Migula. 
(Syst. d. Bakt., 2, 1900, 168.) Hucker 



FAMILY MICKOCOCCACEAE 



97; 



(loc. cit., 27) considers this identical witli 
Micrococcus cinnabareus Fliigge. 

Micrococcus scariosus Migula. (Sie- 
bert, Inaug. Diss., Wiirzburg, No. II, 
1894, 9; Migula, Syst. d. Bakt., 2, 1900, 
91.) From a hairbrush. Winslow and 
Winslow {loc. cit., 199) state that this 
appears to be a synonym of Micrococcus 
albus Schroeter. 

Micrococcus scarlatiriosus Trevisan. 
(Trevisan, Batteri itaiiani, 1879, 19; 
Streptococcus rubiginosus Edington, Brit. 
Med. Jour., 1, 1887, 1265; Perroncitoa 
scarlatinosa Trevisan, I generi e le 
specie delle Batteriacee, Milan, 1889, 
29.) From a scarlet fever patient. 

Micrococcus scarlatinus ^ligula. 
(Syst. d. Bakt., 2, 1900, 173.) From 
feces. 

Micrococcus selenicus Brenner. 
(Jahrb. f. wissensch. Botan., 57, 1916, 85; 
Abst. in Cent. f. Bakt., II Abt., 48, 1918, 
431.) From mud. 

Micrococcus sensibilis Zettnow. 
(Cent. f. Bakt., I Abt., Orig., 77, 1915, 
216.) From dust. Hucker (?oc. cz'i., 19) 
considers this a synonj'm of Micrococcus 
freudenreichii Guillebeau or of Micrococ- 
cus ureae Cohn. For a description of 
this species, see Bergey et al., Manual, 
5th ed., 1939, 248. 

Micrococcus septicus (Klebs) Cohn. 
{Microsporon septicum Klebs, Die Ur- 
sachen der infectiosen Wundkrankheiten, 
1871; and Zur path. Anat. d. Schuss- 
wunden, 1872; Cohn, Beitr. z. Biol. d. 
Pflanzen, 1, Heft 2, 1872, 164.) From 
pus. 

Micrococcus serophilus Costa. 

(Compt. rend. Soc. Biol., Paris, 83, 1920, 
931.) From acute articular rheumatism. 

Micrococcus serratus ^Migula. (Xo. 
15, Lembke, Arch. f. Hyg., 26, 1896, 309; 
Migula, Syst. d. Bakt., 2, 1900, 200.) 
From feces. Winslow and Winslow {loc. 
cit., 205) regard this as a synonym of 
Micrococcus candidus Cohn or of Gaffkya 
tetragena Trevisan. 

Micrococcus sialosepticus Trevisan. 
(Coccwssa^u'ariMS sep<iCMsBiondi,Ztschr. 



f. Hyg., 2, 1887, 195; Coccus septicus 
Biondi, ibid., 220; Trevisan, I generi e 
le specie delle Batteriacee, ^Milan, 1889, 
33; Micrococcus salivarius Migula, Syst. 
d. Bakt., 2, 1900, 65; Micrococcus sali- 
varius -septicus Chester, Man. Determ. 
Bact., 1901, 87.) From human saliva. 
Winslow and Winslow {loc. cit., 205) con- 
sider this a synonym of Micrococcus can- 
didus Cohn or of Gaffkya tetragena Tre- 
visan. 

Micrococcus siccus Migula. (Micro- 
coccus Xo. V, Adametz, Landwirtsch. 
Jahrb., 18, 1889, 241; Migula, Syst. d. 
Bakt., 2, 1900, 124.) From Emmenthal 
cheese. Winslow and Winslow {loc. cit., 
185) state that this is probably a synonym 
of Micrococcus aurantiacus Cohn ; while 
Hucker {loc. cit., 7) considers it a syn- 
onym of Micrococcus luteus Cohn or of 
Micrococcus varians Migula. 

Micrococcus similis Dyar. (Ann. N. 
Y. Acad. Sci., 8, 1895, 347.) From dust. 
Winslow and Winslow {loc. cit., 205) re- 
gard this as a synonym of Micrococcus 
candidus Cohn or of Gaffkya tetragena 
Trevisan. 

Micrococcus simplex Wright. (Mem. 
Xat. Acad. Sci., 7, 1895, 432.) From 
Schuylkill River water. Winslow and 
Winslow {loc. cit., 199) state that this 
appears to be a synonym of Micrococcus 
albus Schroeter. 

Micrococcus simulans DeToni and 
Trevisan. {Micrococcus citreus II, Mag- 
giora, Giorn. Soc. Ital. d'Igiene, 11, 1889, 
354; DeToni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1079.) 

Micrococcus sordidus Schroeter. 
(Schroeter in Cohn, Kryptogam. -Flora 
V. Schlesien, 3, 1, 1886, 145.) Winslow 
and Winslow {loc. cit., 224) state that 
this is apparently a synonym of Micrococ- 
cus candicans Fliigge. 

Micrococcus sphaeroides Gray and 
Thornton. (Cent. f. Bakt., II Abt., 73, 
1928, 74.) From manure and soil. For 
a description of this species, see Bergey 
et al., Manual, 5th ed., 1939, 259. 

Micrococcus staphylophagus Serbinov. 



276 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



(La Defense des Plant es, Leningrad, 2, 
1926, 556; see Rev. AppL MycoL, 5, 1926, 
650.) Considered pathogenic on grape- 
vines. 

Micrococcus stellatus (Lustig) Frank- 
land and Frankland. (Stern-Coccus, 
Maschek, Jahresber. d. Kom.- Oberreal- 
schule, Leitmeritz, Xo. 10, 1887, 62; 
Coccus stellaiiis Lustig, Diag. d. Bakt. d. 
Wassers, 2 Aufl., 1893, 40; Frankland and 
Frankland, Micro-organisms in Water, 
1894, 503.) From water. Winsolw and 
Winslow (loc. cit., 220) regard this as a 
synonym of Micrococcus lutevs Cohn. 

Micrococcus strobiliforrnis Migula. 
(No. 23, Lembke, Arch. f. Hyg., 26, 1896, 
315; Migula, Syst. d. Bakt., 2, 1900, 203.) 
From feces. Winslow and Winslow (loc. 
cit., 220) regard this as a synonym of 
Micrococcus luteus Cohn. 

Micrococcus svbcandicans Lavanchy. 
(Univ. Geneve, Inst. hot. Prof. Chodat, 
Ser. 8, Fasc. 12, 1914, 68; Abst. in Cent, 
f. Bakt., II Abt., 47, 1917, 611.) From 
water of Lake Geneva. 

Micrococcus subcanus Migula. (No. 
17, Lembke, Arch, f . Hyg., 26, 1896, 311 ; 
Migula, Syst. d. Bakt., 2, 1900, 202.) 
From feces. Winslow and Winslow (loc. 
cit., 224) state that this is apparently a 
synonym of Micrococcus candicans 
Fliigge. 

Micrococcus subcarneus Migula. {Mi- 
crococcus carnicolor Kern, Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 495; not 
Micrococcus carnicolor Frankland and 
Frankland, Micro-organisms in water, 
1894, 503; Migula, Syst. d. Bakt., 2, 
1900, 181.) From the intestines of doves 
{Columba livia and Columba oenas). 
Hucker {loc. cit., 26) states that this may 
be identical with Micrococcus roseus 
Fliigge. 

Micrococcus subcitreus Migula. 
(Citronengelber Micrococcus, Keck, 
Ueber das Verhalten der Bakterien im 
Grundwasser, Dorpat Dissertation, 1890, 
60; Migula, Syst. d. Bakt., 2. 1900, 147.) 
From air and water. Winslow and Wins- 
low {loc. cit., 216) consider this a syn- 



onym of Micrococcus flavus Trevisan. 
For a description of this species, see 
Bergey et al.. Manual, 5th ed., 1939, 249. 

Micrococcus subcretaceus Migula. 
(Kreideweisserverfllissigender Micrococ- 
cus, Keck, Inaug. Diss., Dorpat, 1890, 
64; Migula, Syst. d. Bakt., 2, 1900, 107.) 
Winslow and Winslow {loc. cit., 199) 
state that this species appears to be a 
synonym of Micrococcus albus Schroeter. 

Micrococcus subflavescens Bergey et al. 
(Manual, 1st ed., 1923, 61.) From dust 
and water. Hucker {loc. cit., 9) con- 
siders this a synonym of Micrococcus 
flavus Trevisan. For a description of 
this species, see Bergey et al., Manual, 
5th ed., 1939, 246. 

Micrococcus subflavidus Migula. {Mi- 
crococcus tetragenus subflavus v. Besser, 
Beitr. z. allgm. Path. u. path. Anat., 6, 
1889, 347; Migula, Syst. d. Bakt., 2, 1900, 
190; Micrococcus subflavus Chester, Man. 
Determ. Bact., 1901, 96; not Micrococcus 
subflavus Fliigge, Die Mikroorganismen, 
2 Aufl., 1886, 159.) From normal nasal 
mucus. Winslow and Winslow {loc. cit.. 
184) state that this is apparently a syn- 
onym of Micrococcus aureus Zopf ; while 
Hucker {loc. cit., 7 and 21) considers it 
probably identical with Micrococcus lu- 
teus Cohn, Micrococcus varians Migula, 
or Gaffkya tetragena Trevisan. 

Micrococcus subflavus Fliigge. (Gelb- 
weisser Diplococcus, Bumm, Die Mikro- 
org. (1. gonorrh. Schleimhauterkr., 1 
Aufl., 1885 and 2 Aufl., 1887, 20; Fliigge, 
Die Mikroorganismen, 2 Aufl., 1886, 159; 
Neisseria subflava Trevisan, I generi e 
le specie delle Batteriacee, Milan, 1889, 
32; Diplococcus subflavus Eisenberg, 
Bakt. Diag., 3 Aufl., 1891, 307; not 
Micrococcus subflavus Chester, Man. De- 
term. Bact., 1901, 96.) From gonorrheal 
pus. Winslow and Winslow {loc. cit., 
216) consider this a synonym of Mi- 
crococcus flavus Trevisan. For a descrip- 
tion of this species, see Bergey et al., 
Manual, 5th ed., 1939, 248. 

Micrococcus subftisciis Matzuschita. 
(Cent. f. Bakt., I Abt., 29, 1901, 383.) 



FAMILY MICROCOCCACEAE 



277 



From dust. Similar to Micrococcus Jus- 
ens Adametz. 

Micrococcus suhgilvus Migula. {Mi- 
crococcus gilvus Henrici, Arb. a bakt. 
Inst. Karlsruhe, 1, Heft 1, 1894, 78; not 
Micrococcus gilvus Losski, Inaug. Diss., 
Dorpat, 1893, 60; Migula, Syst. d. Bakt., 
2, 1900, 132.) From cheese. Winslow 
and Winslow {loc. cit., 220) regard this 
as a synonj'^m of Micrococcus luteus 
Cohn. 

Micrococcus subgranulatus INIigula. 
(Micrococcus citreus granulatus Freund, 
Inaug. Diss., Erlangen, 1893, 27; Migula, 
Syst. d. Bakt., 2, 1900, 148; Micrococcus 
granulatus v. Bagarewski, Cent, f . Bakt., 
II Abt., 15, 1905, 7.) From the oral 
cavity. Winslow and Winslow {loc. cit., 
216) consider this a sj'nonjon of Micro- 
coccus flavus Trevisan. For a descrip- 
tion of this species, see Bergej^ et al., 
Manual, 5th ed., 1939, 249. 

Micrococcus subgrise^ts Migulq., 
(Grauer Coccus, Maschek, Jahresb. d. 
Kom.- Oberrealschule, Leitmeritz, No. 
8, 1887, 61; Migula, Syst. d. Bakt., 2, 
1900, 94.) From water. Winslov/ and 
Winslow (loc. cit., 199) state tliat this 
appears to be a synonym of Micrococcus 
albus Schroeter; while Hucker (loc. cit., 
19) regards it as a synonym of Micrococcus 
freudenreichii Guillebeau or of Micro- 
coccus ureae Cohn. 

Micrococcus sublacteus Migula. (No. 
27, Lembke, Arch. f. Hj'g., 29, 1897, 329; 
Migula, Syst. d. Bakt., 2, 1900, 210.) 
From feces. Winslow and Winslow (loc. 
cit., 199) state that this appears to be a 
S3-nonym of Micrococcus albus Schroeter; 
while Hucker (loc. cit., 19) regards it as 
a synonym of Micrococcus freudenreichii 
Guillebeau or of Micrococcus ureae Cohn. 

Micrococcus sublilacinus Migula. (No. 
26, Lembke, Arch. f. Hyg., 26, 1896, 317; 
Migula, Syst. d. Bakt., 2, 1900, 205.) 
From feces. Hucker (loc. cit., 15) con- 
siders this a synonym of Micrococcus al- 
bus Schroeter. 

Micrococcus subluteus Weiss. (Arb. 



bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
198.) From vegetable infusions. 

Micrococcus subniveus Migula. (Mi- 
crococcus albidus Henrici, Arb. bakt. 
Inst. Karlsruhe, 1, Heft 1, 1894, 75; not 
Micrococcus albidus Losski, Inaug. Diss., 
Dorpat, 1893, 55; Migula, Syst. d. Bakt., 
2, 1900, 105.) From Swiss cheese. 
Winslow and Winslow (loc. cit., 199) state 
that this appears to be a synonym of 
Micrococcus albus Schroeter. 

Micrococcus subochraceus Migula. 
(No. 30, Lembke, Arch. f. Hyg., 29, 1897, 
332; Migula, Syst. d. Bakt., 2, 1900, 215.) 
From feces. Winslow and Winslow (loc. 
cit., 216) regard this as a synonym of 
Micrococcus flavus Trevisan. 

Micrococcus subroseus Migula. (Mi- 
crococcus roseus Eisenberg, Bakt. Diag., 
3 Aufl., 1891, 408; Migula, Syst. d. Bakt., 
2, 1900, 176 ; Micrococcus rubescens Ches- 
ter, Man. Determ. Bact., 1901, 105.) 
From the sputum of an influenza patient. 
Hucker (loc. cit., 26) states that this may 
be identical with Micrococcus roseus 
Flligge. 

Micrococcus subterraneus Hansgirg. 
(Hansgirg, Oest. Bot. Zeitschr., 1888, 
No. 7-8, 8; Staphylococcus subterraneus 
DeToni and Trevisan in Saccardo, 
Sylloge Fungorum, 8, 1889, 1075.) From 
damp walls of wine cellars in Bohemia. 

Micrococcus subtilis Migula. (Diplo- 
coccus, Kirchner, Ztschr. f . Hyg., 9, 1890, 
528; Migula, Syst. d. Bakt., 2, 1900, 192.) 
Found in the sputum and Mood of influ- 
enza patients. 

Micrococcus succulentus Henrici. 
(Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 
1894, 63.) From Swiss cheese. Wins- 
low and Winslow (loc. cit., 224) state 
that this is apparently a synonym of 
Micrococcus candicans Flligge. 

Micrococcus sulphur eus Zinamermann. 
(Bakt. unserer Trink- u. Nutzwasser, 
Chemnitz, I Reihe, 1890, 84.) From 
water. Winslow and Winslow (loc. cit., 
220) regard this as a synonym of Micro- 
coccus luteus Cohn. 

Micrococcus suis Burrill. (Bacillus 



278 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



stiis Detmers, Rept. U. S. Dept. Agric. 
for 1878; Burrill, Amer. Nat., 17, 1883, 
320.) From blood of hogs sick with 
swine plague or hog cholera. 

Micrococcus syphiliticus Migula. (Coc- 
cen, Disse, Deutsche med. Wchnschr., 
13, 1887, 888; Migula, Syst. d. Bakt., 2, 

1900, 218.) This may be synonymous 
with Micrococcus candicans Fliigge. 

Micrococcus tardigradus Trevisan. 
{Micrococcus flavus tardigradus Fliigge, 
Die Mikroorganismen, 2 Aufi., 1886, 175; 
Trevisan, I generi e le specie delle Bat- 
teriacee, Milan, 1889, 34; Micrococcus 
sulfureus fi-tardigradus Lehmann and 
Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 
163; Micrococcus sulfureus var. tar- 
digradus Lohnis and Pillai, Cent. f. 
Bakt., II Abt., 19, 1907, 92.) From air; 
also found in water. Winslow and Wins- 
low {loc. cit., 220) regard this as a 
synonym of Micrococcus luteus Cohn. 

Micrococcus tardior Migula. {Diplo- 
coccus flavus liquefaciens tardus Unna 
and Tommasoli, Monatshefte f. prakt. 
Dermatol., 9, 1889, 56; Migula, Syst. d. 
Bakt., 2, 1900, V and 141; Micrococcus 
epidermis Chester, Man. Determ. Bact., 

1901, 97; Diplococcus flavus -lique J aciens 
Chester, ibid.) From eczema. Wins- 
low and Winslow {loc. cit., 216) regard 
this as a synonym of Micrococcus flavus 
Trevisan; while Hucker {loc. cit., 11) 
regards it as a synonym of Micrococcus 
citreus Migula. 

Micrococcus tardissimus (Trevisan) 
Migula. (Milchweisser Micrococcus, 

Bumm, Mikroorg. d. gonorrh. Schleim- 
hauterkr., 1 Aufl., 1885; Diplococcus 
albicans tardissimus Fliigge, Die Mikro- 
organismen, 2 Aufl., 1886, 183; Neisseria 
tardissima Trevisan, I generi e le specie 
delle Batteriacee, Milan, 1889, ^2; Micro- 
coccus albicans tardissimus Sternberg, 
Man. of Bact., 1893, 882; Migula, Syst. 
d. Bakt., 2, 1900, 49.) Found in vaginal 
secretions. Winslow and Winslow {loc. 
cit., 205) regard this as a synonym of 
Micrococcus candidus Cohn or of Gaffkya 
tetragena Trevisan; while Hucker {loc. 



cit., 7) considers it a synonym of Micro- 
coccus luteus Cohn or Micrococcus varians 
Migula. 

Micrococcus tardus Migula. {Diplo- 
coccus albicans tardus Unna and Tom- 
masoli, Monatshefte f. prakt. Dermatol., 
9, No. 2, 1889, 49; Micrococcus albicans 
tardus Sternberg, Man. of Bact., 1893, 
882; Migula, Syst. d. Bakt., ^, 1900, 50; 
Micrococcus eczemae Chester, Man. De- 
term. Bact., 1901, 86). From eczema. 
Winslow and Winslow {loc. cit., 216 and 
224) regard this as a synonym of Micro- 
coccus flavus Trevisan or of Micrococcus 
candicans Fliigge. 

Micrococcus tenacatis Chester. (No. 
43, Conn, Storrs Agr. Exp. Sta. 7th Ann. 
Rept., 1895, 78; Chester, Man. Determ. 
Bact., 1901, 88.) From milk from Uru- 
guay. Winslow and Winslow {loc. cit., 
220) state that this is apparently a syn- 
onym of Micrococcus candicans Fliigge. 

Micrococcus tener Weiss. (Arb. bakt. 
inst. Karlsruhe, 2, Heft 3, 1902, 200.) 
From a vegetable infusion. 

Micrococcus ienMzssun?/s Migula. {Mi- 
crococcus cumulatus tenuis v. Besser, 
Beitr. z. path. .\nat.,6, 1889, 347 ; Migula, 
Syst. d. Bakt., 2, 1900, 55; Micrococcus 
cumulatus Chester, Man. Determ. Bact., 
1901, 87.) Frequently found in human 
nasal mucus. Winslow and Winslow 
{loc. cit., 205) regard this as a synonym 
of Micrococcus candidus Cohn or of 
Gaffkya tetragena Trevisan. 

Micrococcus tetragenus aureus Boutron. 
(These, Paris, 1893; Abst. in Cent. f. 
Bakt., 16, 1894, 971.) Hucker {loc. cit., 
21 ) regards this as a synonym of Gaffkya 
tetragena Trevisan. 

Micrococcus tetragenus concentricus 
Schenk. (Allg.Wien. med. Zeitung, 1892, 
81 and 92; Abst. in Cent. f. Bakt., 13, 
1893, 720.) From feces. Motile. 

Micrococcus tetragenus -pallidus Ches- 
ter. {Micrococcus tetragenus pallidus, 
Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 354; 
Chester, Man. Determ. Bact., 1901, 93.) 
From dust. Probably a variety of Mi- 
crococcus versatilis Chester, see below. 



FAMILY MICROCOCCACEAE 



279 



Micrococcus tetragenvs-vividiis Ches- 
ter. (Micrococcus tetragemts vividus 
Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 354 ; 
Chester, Man. Determ. Bact., 1901, 102.) 
From dust . Probably a variety of Micro- 
coccus versaiilis Chester, see below. 

Micrococcus tetras Henrici. (Arb. 
bakt. Inst. Karlsruhe, /, Heft 1, 1894, 
60; Pediococcus tetras Pribram, Klassi- 
fikation der Schizomyceten, 1933, 46.) 
From cheese. Winslow and Winslow 
(loc. cit., 224) state that this species is 
apparently the same as Micrococcus 
candicans Fliigge. 

Micrococcus thermophilus Hansgirg. 
(Oestr. Bot. Ztschr., No. 3, 1888, 5.) 
From hot springs. 

Micrococcus toxicatus Burrill. (Amer. 
Xat., 17, 1883, 319.) From poison ivy 
and other plants in the genus Rhus. 

Micrococcus trachomatis IMigula. 
(Trachomococcus, Sattler, in Zehender, 
Klin. Monatsbl., 1881; Trachomococcus, 
Michel, Arch. f. Augenheilk., 16, 1886; 
Neisseria micheli Trevisan, I generi e 
le specie delle Batteriacee, Milan, 1889, 
32; see Baumgarten, Lehrb. d. path. 
Mykol., 1, 1890, 421; Migula, Syst. d. 
Bakt., 2, 1900, 67.) Winslow and Wins- 
low (loc. cit., 205) consider this to be a 
synonym of Micrococcus candidus Cohn 
or of Gaffkya tetragena Trevisan. 

Micrococcus trihutyrus Stark and 
Scheib. (Jour. Dairy Sci., 19, 1936, 210.) 
From butter. 

Micrococcus tritici Prillieux. (Mala- 
dies des plantes agricoles, 1, 1895, 7; not 
Micrococcus tritici Kock, Monatshefte f. 
Landwirtschaft, 1909, 247, quoted from 
Lehmann and Neumann, Bakt. Diag., 
5 Aufl., 2, 1912, 653.) Considered patho- 
genic on wheat. 

Micrococcus tuberculosus Migula. 
(No. 23, Lembke, Arch. f. Hyg., 29, 1897, 
325; Migula, Syst. d. Bakt., 2, 1900, 214.) 
From feces. Winslow and Win.slow (loc. 
cit., 224) state that this is apparently a 
synonym of Micrococcus candicans Fliigge. 



Micrococcus typhoideus Migula. (Coc- 
cus A, Foutin, Bakt. Untersuch. v. Hagel, 
Wratsch, 1889, No. 49 and 50; see Cent, 
f. Bakt., 7, 1890, 373; Migula, Syst. d. 
Bakt., 2 1900, 94; Micrococcus alpha 
Chester, Man. Determ. Bact., 1901, 93.) 
From hail. Winslow and Winslow (loc. 
cit., 199) state that this appears to be a 
synonym of Micrococcus alb^is Schroeter ; 
while Hucker (loc. cit., 25) states that it 
may be identical with Micrococcus roseus 
Fliigge. 

Micrococcus ulceris de Luca. (Gaz- 
zetta degli Ospitali, 1886; Abst. in Cent, 
f. Bakt., 1, 1887, 333; Micrococcus ulceris 
mollis de Luca, ibid.) From the secre- 
tion of a venereal ulcer. 

Micrococcus ulmi Brussoff. (Cent. f. 
Bakt., II Abt., 63, 1925, 261.) Isolated 
from diseased elm trees. 

Micrococcus umbilicatus Weiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
186.) From a bean infusion. Hucker 
(loc. cit., 12) considers this a s.ynonym of 
Micrococcus aureus Zopf. 

Micrococcus urcae Migula. (Torule 
ammoniacale, Pasteur, Ann. de Chim. 
et de Phys., 3 ser., 64, 1862, 52; van 
Tieghem, Comp. rend. Acad. Sci., Paris, 
58, 1864, 210; Torula ureae Lea, Jour, of 
Physiol., 11, 1890, 226; Migula, in Engler 
and Prantl, Die natiirl. Pflanzenfam., 
/, la. 1895, 17.) From urine. May not 
be the same as Micrococcus ureae Cohn. 

Micrococcus urinalbus De Toni and 
Trevisan. [Micrococcus albus urinae 
Doyen, Jour. d. connaiss. medic, No. 
14, 1889, 108; De Toni and Trevisan, in 
Saccardo, S3dIoge Fungorum, 8, 1889, 
1076.) From urine. Hucker {loc. cit., 
15) considers this a synonym of Micrococ- 
cus albus Schroeter. 

Micrococcus uruguae Chester. (No. 
40, Conn, Storrs Agr. Exp. Sta. 7th Ann. 
Kept., 1895, 78; Chester, Man. Determ. 
Bact., 1901, 100.) From milk from 
Uruguay. Winslow and Winslow (loc. 
cit., 216) regard this as a synonym of 



280 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Micrococcus flavus Trevisan; while 
Hucker {loc. cit., 10) regards it as a 
synonym of Micrococcus conglomeratus 
Migula. 

Micrococcus utriculosiis Migula. (No. 
20, Lembke, Arch. f. Hyg., 29, 1897, 327; 
Migula, Syst. d. Bakt., 2, 1900, 199.) 
From feces. Winslow and Winslow {loc. 
cit., 199) state that this appears to be a 
synonym of Micrococcus alhus Schroeter. 

Micrococcus varians lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900, 37.) From milk, cream, dust. 
According to Weigmann (In Lafar, 
Handb. d. techu. Mykologie, 2, 1905, 
13) this is identical with Staphylococcus 
mastitis albus. Hucker {loc. cit., 11) 
regards it as a synonym of Micrococcus 
citreiis Migula. 

Micrococcus variococcus Miiller-Thur- 
gau and Osterwalder. (Cent. f. Bakt., 
II Abt., 33, 1913, 23G.) From wine. 

Micrococcus versatilis Chester. {Mi- 
crococcus tetragenus febris flavae Finlaj^; 
Micrococcus tetragenus versatilis Stern- 
berg, Report on etiology and prevention 
of yellow fever, Washington, 1891, 164; 
Chester, Man. Determ. Bact., 1901, 102.) 
Isolated from the excrement of mosqui- 
toes which had sucked the blood of yellow 
fever patients; and from dust. Winslow 
and Winslow {loc. cit. ,21Q) regard this as a 
synonym of Micrococcus flavus Trevisan. 

Micrococcus versicolor Flligge. (Die 
Mikroorganismen, 2 Aufl., 1886, 177.) 
From dust. Winslow and Winslow {loc. 
cit., 220) consider this a synonym of 
Micrococcus luteus Cohn. 

Micrococcus vesicae Heim. (Lehrb. 
d. Bakt., 2 Aufl., 1898, 297.) From acid 
urine. Winslow and Winslow {loc. cit., 
224) state that this is apparently a 
synonym of Micrococcus candicans Fliigge. 
Micrococcus vesicans Harman. (Jour. 
Path, and Bact., 9, 1904, 1.) Considered 
the cause of veld sore, a disease of Africa 
and tropical Australia. 

Micrococcus vesicas us Weiss. (Arb. 
bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 
203.) From a vegetable infusion. 



Hucker {loc. cit., 8) considers this spe- 
cies identical with either Micrococcus 
luteus Cohn or Micrococcus varians 
Migula. 

Micrococcus vesiculiferus Migula. 
(No. 28, Lembke, Arch. f. Hyg., 29, 1897, 
330; Migula, Syst. d. Bakt., 2, 1900, 211.) 
From feces. Winslow and Winslow {loc. 
cit., 220) regard this as a synonym of 
Micrococcus luteus Cohn. 

Micrococcus vincenzii Chester. {Mi- 
crococcus tetragenus citreus Vincenzi, La 
Riforma Med., 1897, 758; Chester, Man. 
Determ. Bact., 1901, 103.) From the 
submaxillary lymphatic gland of a child. 
Winslow and Winslow {loc. cit., 220) re- 
gard this as a synonym of Micrococcus 
luteus Cohn. 

Micrococcus vini Migula. {Micrococ- 
cus saprogenes vini I, Kramer, Bakt. in 
Beziehungen z. Landwirtsch. u. d. land- 
wirtsch-techn. Gewerben, II Teil, 1892, 
139; Migula, Syst. d. Bakt., 2, 1900, 118.) 
From wine. Winslow and Winslow {loc. 
cit., 199) state that this appears to be a 
synonym of Micrococcus albus Schroeter; 
while Hucker {loc. cit., 8) considers it 
identical with Micrococcus luteus Cohn 
or Micrococcus varians Migula. 

Micrococcus viniperda Schroeter. 
(Schroeter in Cohn, Kryptog. -Flora v. 
Schlesien, 3, 1, 1886, 144.) From dust, 
feces, etc. 

Micrococcus viscosus Bergey et al. 
{Micrococcus lactis viscosus B, Conn, 
Esten and Stocking, Storrs Agr. Exp. 
Sta. Rept. for 1906, 109; Bergey et al., 
iManual, 1st cd., 1923, 68.) From pas- 
teurized milk. For a description of this 
species, see Bergey et al., Manual, 5th 
ed., 1939, 256. See Micrococcus lactis 
viscosus Sternberg. 

Micrococcus viscosus lactis Conn. 
(Storrs Agr. Exp. Sta. 12th Ann. Rept., 
1900, 44.) From milk. 

Micrococcus viticulosus Fliigge. (Die 
Mikroorganismen, 2 Aufl., 1886, 178.) 
From dust and water. Winslow and 
Winslow {loc. cit., 205) consider this to 



FAMILY MICROCOCCACEAE 



281 



be a synonym of Micrococcus candidiis 
Cohn or of Gaffkya tetragena Trevisan. 

Micrococcus vulgaris Eckstein. 
(Ztschr. f. Forst- u. Jagdwesen, 26, 
1894, 17; Abst. in Cent. f. Bakt., I Abt., 
18, 1895, 292; not Micrococcus vulgaris 
Weiss, Arb. bakt. Inst. Ivarlsruhe, 2, 
Heft 3, 1902, 193.) From insects. 

Micrococcus xanthogenicxis (Freire) 
Trevisan. {Cryptococcus xanthogenicus 
Freire, Recherches sur la cause de la 
fievre jaune, Rio de Janeiro, 1884; Tre- 
visan, I generi e le specie delle Batteria- 
cee, Milan, 1889, 33.) Isolated from 
yellow fever and supposed by Freire to 
be the cause of the disease. Winslow 
and Winslow {loc. cit., 199) state that 
this appears to be a synonym of Microco- 
cus albus Schroeter. 

Micrococcus xenopus Schrire and Green- 
field. (Trans. Royal Soc. So. Africa, 
17, 1930, 309.) From an abscess in a toad 
(Xenopus sp.). For a description of 
this species, see Bergcy et al.. Manual, 
5th ed., 1939, 243. 

Micrococcus xerophilus Glage. 

(Ztschr. f. Fleisch- u. IMilchhygiene, 10, 
1900, 145.) From coating on surface of 
dry wurst and similar meat products. 

Micrococcus zeae Serbinov. (La De- 
fense des Plantes, S, 1926, 546.) From 
flour, grain and seedlings of corn. Was 
thought to be a cause of pellagra in South 
Russia. 

Micrococcus zonatus Henrici. (Arb. 
bakt. Inst. Karlsruhe, i, Heft 1, 1894, 68.) 
From cheese. Winslow and Winslow 
(loc. cit., 224) state that this is appar- 
ently a synon}^m of Micrococcus candi- 
cans Fliigge. 

Planococcus casei iNIigula. (ISIicro- 
coccus No. Ill, Adametz, Landwirtsch. 
Jahrb., 18, 1889, 240; Migula, Syst. d. 
Bakt., 2, 1900, 270.) From Emmenthal 
cheese. 

F lanococcus loeffleri Migula. (Loffler, 
Cent. f. Bakt., 7, 1890, 637; Migula, Syst. 
d. Bakt., 2, 1900, 273.) From colony on 
an old gelatin plate. 

Planococcus luteus (Adametz) Migula. 



(Diplococcus luteus Adametz, Mitteil. 
d. oesterr. Vers. Station f. Brauerei u. 
Malzerei in Wien, Heft I, 1888, 39; 
Neisseria lutea Trevisan, I generi e le 
specie delle Batteriacee, Milan, 1889, 32; 
Migula, Syst. d. Bakt., 2, 1900, 274.) 
Hucker {loc. cit., 9) considers this species 
a synonym of Micrococcus fiavus Trevi- 
san. 

Rhodococcus fulvus Winslow and 
Rogers. (Jour. Inf. Dis., 8, 1906, 545; 
not Micrococcus fulvus Cohn, Beitr. z. 
Biol. d. Pflanzen, 1, Heft 3, 1875, 181.) 
From soil, air and water. 

Staphylococcus albicans Stigell. 
(Cent. f. Bakt., I Abt., Orig., 45, 1908, 
489.) Probably intended for Micrococcus 
albicans amplus Fliigge. 

Staphylococcus albus liquefaciens 
Sternberg. (White liquefying staphjdo- 
coccus, Escherich, Die Darmbakterien 
des Sauglings, Stuttgart, 1886, 88; Stern- 
berg, Manual of Bact., 1893, 607.) Found 
occasionalh^ in the feces of healthy 
infants. 

Staphylococcus albus non liquefaciens 
Hlava. (Sbornik lekafsky, II, Prague, 
1887, 12 pp.; see Cent. f. Bakt., 2, 1887, 
688.) Probably a synonym of Micrococ- 
cus albocereus INIigula. 

Staphylococcus anaerobius Heurlin. 
(Bakt. Unters. d. Keimgehaltes im 
Genitalkanale d. fiebernden Wochnerin- 
nen, Helsingfors, 1910, 120.) See 
Weinberg, Xativelle and Prevot, Les 
INIicrobes Anaerobies, Paris, 1937, 1027; 
probably not the same as Staphylococcus 
anaerobius Hamm, Die puerperale 
Wundinfektion, Berlin, 1912. Incom- 
pletely described. From genital tract. 

Staphylococcus anaerobius major Heur- 
lin {loc. cit., 120). From genital tract. 

Staphylococcus anaerobius minor Heur- 
lin {loc. cit., 120). From genital tract. 

Staphylococcus aureus sarciniforinis 
Rosenhauch. (Klin. Monatsbl. f . Augen- 
heilkunde, Jahrg., 8, 1909, 257; Abst. in 
Cent. f. Bakt., I Abt., Ref., 45, 1910,787.) 

Staphylococcus bovis Ford. {Staphylo- 
coccus pyogenes boi-is Lucet, Ann. Inst. 



282 



MANUAL 0F DETERMINATIVE BACTERIOLOGY 



Past., 7, 1893, 327 ; Ford, Textb. of Bact., 
1927, 424.) Found in suppurative le- 
sions of cattle. 

Staphylococcus candidus Warrington. 
(Lancet, 1, 1888.) 

Staphylococcus flavocyaneus Knaysi. 
(Jour. Bact., 43, 1942, 368.) Found as a 
contaminant in dissociation studies. 

Staphylococcus flavus nan pyogenes 
Frankel and Sanger. (Arch. f. patli. 
Anat., 108, 1887, 286; Abst. in Cent. f. 
Bakt., 3, 1888, 281.) Found in endo- 
carditis ulcerosa. 

Staphylococcus griseus Tavel. (Quoted 
from Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., 2, 1896, 173.) From pus. 

Staphylococciis griseus radiatus Viti. 
(Atti d. R. Accad. d. Fisiocritici di 
Siena, Ser. IV., 2, 1891, Abst. in Cent, 
f. Bakt., 11, 1892, 672.) From cases of 
endocarditis. 

Staphylococcus habanensis Gibler. 
(Quoted from Fernandez, Cronica medico 
quiriirgica de la Habana, 1891, No. 30; 
Abst. in Cent. f. Bakt., 11, 1892, 472.) 
Isolated from the human eye. 

Staphylococcus insectorum Krassilst- 
schik. (Quoted from Paillot, Les mal- 
adies du ver a sole grasserie et dysen- 
teries, 1928, 171.) From the intestinal 
tract of the silkworm {Bombyx mori). 

Staphylococcus lactis acidi McDonnell. 
(Inaug. Diss., Kiel, 1899; Abst. in Cent, 
f. Bakt., II Abt., 6, 1900, 120.) 

Staphylococcus leloirii Trevisan. (Mi- 
crobe des perifolliculites conglomerees, 
Leloir, Soc. anatomique, May, 1884; Tre- 
visan, I generi e le specie delle Batteria- 
cee, Milan, 1889,33.) 

Staphylococcus liqucfaciens aurantia- 
cus Distaso. (Cent. f. Bakt., I Abt., 
Orig., 59, 1911, 102.) From feces. 

Staphylococcus muscae Glaser. (Amer. 
Jour. Hyg., 4, 1924, 411.) Causes a fatal 
infection in house flies {M usca dornestica) . 
For a description of this species, see 
Bergey et al.. Manual, 5th ed., 1939, 264. 

Staphylococcus non pyogenes Savor. 
(Beitr. z. Geburtshilfe u. Gynakol. v. 
Hegar, 2, Heft 1, 1898; Abst. in Cent. f. 



Bakt., I Abt., 26, 1899, 642.) From 
urino-genital tract. 

Staphylococcus pharyngis Bergey et al. 
(Manual, 1st ed., 1923, 56.) Found in 
the human nasopharynx in acute catarrh- 
al inflammation. For a description of 
this species, see Bergey et al.. Manual, 
5th ed., 1939, 265. 

Staphylococcus putrificus Schottmuller. 
(Leitfaden f. d. Klinisch- Bakt. Kultur- 
Methoden, Berlin, Wien, 1923. Quoted 
from Weinberg, Nativelle and Prevot, 
Les Microbes Anaerobies, Paris, 1927, 
1027.) 

Staphylococcus pyogenes liquefaciens 
albus Hlava. (Sbornik lekafsk;^, II, 
Prague, 1887, 12 pp.; Abst. in Cent. f. 
Bakt., 2, 1887, 688.) From small pox 
pustules. 

Staphylococcus pyogenes tenuis Scheibe. 
(Inaug. Diss., Miinchen, 1889; see Cent, 
f. Bakt., 6, 1889, 186.) From middle ear 
infections. 

Staphylococcus roseus Tavel. (Quoted 
from Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., 2, 1896, 177.) Evidently 
identical with Micrococcus roseus Leh- 
mann and Neumann or Micrococcus 
roseo-fulvus Lehmann and Neumann. 

Staphylococcus salivarius Andrewes 
and Gordon. (35th Ann. Rept. Local 
Govt. Board London, 1905-06, 558.) 
From saliva. Probably Micrococcus can- 
didus Cohn. 

Staphylococcus ureae candidus Lund- 
strom. (Festschr. d. path. -anat. Inst, 
z. Andenken a. d. 250 jiihrige Bestehen d. 
finnland. Univ. Helsingfors, 1890; abst. 
in Cent. f. Bakt., 9, 1891, 672.) From 
urine. Probably Micrococcus ureae 

Cohn. 

Staphylococcus ureae non pyogenes 
Barlow. (Arch. f. Dermat. u. Syph., 
1893; Abst. in Cent. f. Bakt., U, 1893, 
456.) From cases of cystitis. 

Urococcus dowdeswelli ^liquel. (Ann. 
de Micrographie, 5, 1893, 209.) Fer- 
ments urea. 

Urococcus van tieghemi Miquel (Joe. 
cit., 161). Ferments urea. 



FAMILY MICROCOCCACEAE 



283 



Genus II. Gaflfkya Trevisan* 

(Trevisan, Atti d. Accad. Fisio-Medico-Statistica in Alilano, Ser. 4, 3, 1885, 106; 
Tetracoccus Klecki, Cent. f. Bakt., 15, 1894, 360; not Tetracocciis Orla-Jensen, The 
Lactic Acid Bacteria, Mem. Acad. Sci. Danemark, Sec. Sci., 8 ser, 5, 1919, 154; Tetra- 
diplococcus Bartoszewicz and Schwarzwasser, Cent. f. Bakt., II Abt., -21, 1908, 614.) 
Named for Prof. Georg Gaffky, 1850-1918, Berlin. 

Occur in the animal body and in special media as tetrads, while in ordinary culture 
media they occur in pairs and irregular masses. Aerobic to anaerobic. Gram-posi- 
tive. Parasitic organisms. 

The type species is Gaffky a tetragena (Gaffky) Trevisan. 



I. Facultative aerobe. 
II. Strict anaerobe. 



Key to the species of genus Gaffkya. 

1. Gaffkya tetragena. 

2. Gaffkya anaerobia. 



1. Gaffkya tetragena (Gaffky) Trevi- 
san. {Micrococcus tetragenus Gaffky, 
Arch. f. Chirurg., 38, Heft 3, 1883, 500; 
Trevisan, Atti d. Accad. Fisio- Medico- 
Statistica in Milano, Ser. 4, 3, 1885, 106; 
Micrococcus tetragenus septicus Boutron, 
Thesis, Paris, 1893; Abst. in Cent. f. 
Bakt., 16, 1894, 971; Micrococcus tetra- 
genus albus Boutron, ibid.; Merista sep- 
tica Hueppe, Principles of Bacteriology 
(Eng. trans.), 1899, 170; Sarcina seplica 
Hueppe, ibid.; Sarcina tetragena Migula, 
Syst. d. Bakt., 2, 1900, 225; Merista 
tetragena Vuillemin, Ann. Mj^cologie, 
Berlin, 11, 1913, 525; Staphylococcus 
tetragenus Holland. Jour. Bact., 5, 1920, 
224; Tetracoccus septicus Neveu-Lemaire, 
Precis Parasitol. Hum., 5th ed., 1921, 
18; Pediococcus tetragenus Pribram, Klas- 
sifikation der Schizomyceten, 1933, 46.) 
From Greek, tetra (tetara), four; M. L.- 
ge7ies, producing. 

Spheres : 0.6 to 0.8 micron in size, with 
pseudocapsule (in bodj^ fluids) surround- 
ing four of the elements showing tj-pical 
tetrads. Gram -positive. 



Gelatin colonies: Small, 1 to 2 mm. in 
diameter, white convex. 

Gelatin stab : Thick, white surface 
growth. No liquefaction. 

Agar colonies : Circular, white, smooth, 
glistening, entire. Reimann (Jour. Bact., 
31, 1936, 385) has described eleven 
colony form variants for this species. 

Agar slant: White, moist, glistening. 

Broth : Clear, with gray viscous sedi- 
ment. 

Litmus milk: Slightly acid. 

Potato: White, viscid. 

Indole not formed. 

Nitrites not produced from nitrates. 

Starch not hydrolyzed. 

Ammonium salts not utilized. 

Acid from glucose, lactose and 
glycerol. 

No HoS formed. 

Aerobic, facultative. 

Pathogenic for mice and guinea pigs; 
rabbits less susceptible. 

Optimum temperature 37 °C. 

Source: Isolated from sputum in 
tuberculosis; also from air and skin. 

Habitat : Mucous membrane of respira- 
tory tract. 



* Revised by Prof. G. J. Hucker, N. Y 
York, :March, 1943. 



State Experiment Station, Geneva, New 



284 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



2. Gaffkya anaerobia (Choukevitch) 
Prevot. {Tetracoccus anaerobius Chou- 
kevitch, Ann. Inst. Past., 25, 1911, 349; 
Micrococcus tetragenes anaerobius Hamm, 
Die puerperale Wundinfektion, Berlin, 
1912; Prevot, Ann. Sci. Nat., Ser. Bot., 
15, 1933, 203.) From Greek, an, without ; 
aer, air; bios, life. 

Spheres: About 1.0 to 1.5 microns, 
occurring in tetrads, sometimes in groups 
of eight. Gram-positive. 

Gelatin : No liquefaction. 

Deep agar colonies : After 24 to 48 
hours, small, grayish, 2 to 3 mm. in diam- 
eter. Abundant production of gas which 
breaks up the agar. 

Broth: Poor growth. Slight sediment. 

Milk: Unchanged. 

Coagulated proteins not digested. 

Optimum temperature 37°C. No 

growth at 22 °C. 

Non-pathogenic for guinea-pigs or 
rabbits. 

Strict anaerobe. 

Distinctive characters : Prefers acid 
media. 

Source : Isolated from the female gen- 
ital tract; isolated from the large in- 
testine of a horse. 

Habitat : Probably widely distributed 
in natural cavities of man and animals. 

Appendix: The following species have 
been placed in the genus Gaffkya or in 
the genus Tetracoccus. 

Gaffkya archeri Trevisan. (A black 
micrococcus. Archer, Quart. Jour. Mi- 
croscop. Sci., 1874, 321; Trevisan, I 
generi e le specie delle Batteriacee, Milan, 
1889, 27.) 

Gaffkya grandis DeToni and Trevisan. 
(Microcoque des reins et des ulceres 
syphilitiques de la peau. Babes, in Cornil 
and Babes, Les Bacter., 2nd ed., 1886, 
782; DeToni and Trevisan, in Saccardo, 
Sylloge Fungorum, 8, 1889, 1042.) 

Gaffkya mendozae DeToni and Trev- 



isan. {Micrococcus tetragenus and Mi- 
crococcus tetragenus mobilis ventriculi 
Mendoza, Cent. f. Bakt., 6, 1889, 567; 
DeToni and Trevisan, in Saccardo, Syl- 
loge Fungorum, 8, 1889, 1043 ; Planococcus 
tetragenus Migula, Syst. d. Bakt., 2, 1900, 
269; Micrococcus mendozae Chester, Man. 
Detcrra. Bact., 1901, 84; Sarcina tetra- 
gena Winslow and Rogers, Jour. Inf. Dis., 
3, 1906, 545; Planomerista ventricidi 
Vuillemin, Ann. Mycolog., Berlin, 11, 
1913, 525.) Motile. Isolated from the 
contents of the stomach. Hucker (N. 
Y. State Exp. Sta. Tech. Bull. No. 102, 
1924, 21) regards this as a synonym of 
Gaffkya tetragena Trevisan. 

Gaffkya tardissima (Altana) Bergey 
et al. {Tetragenus tardissiynus Altana, 
Cent. f. Bakt., I Abt., Orig., 48, 1909, 
42; Bergey et al.. Manual, 2nd ed., 1925, 
59.) From a natural infection of guinea 
pigs. See Manual, 5th ed., 1939, 269 for a 
description of this species. 

Gaffkya verneti Corbet. (Organism 
No. 21, Denier and Vernet, La Caout- 
chouc, 17, 1920, 10491; Corbet, Quart. 
Jour. Rubber Research Inst., Malaya, 2, 
1930, 143.) From the latex of the Para 
rubber tree {Hevea brasiliensis) . For a 
description of this species, see Manual, 
5th ed., 1939, 269. 

Tetracoccus carneus halophilus Horo- 
witz-Wlassowa. (Cent, f . Bakt., II Abt., 
85, 1932, 16.) Isolated from salted in- 
testines (Wiener skins). 

Tetracoccus casei Orla-Jensen. (The 
Lactic Acid Bacteria, 1919, 80.) From 
cheese. Probably identical with Micro- 
coccus freudenreichii Guillebeau. 

Tetracoccus mastitidis Orla-Jensen {loc. 
cit., 81). From milk of a woman with 
mastitis. Orla-Jensen thinks this is 
identical with the staphylococcus that 
causes mastitis in cows, i.e., Micrococcus 
pyogenes var. aureus Zopf. 

Tetracoccus mycodcrmatus Orla-Jensen 
{loc. cit., 81). From Camembert cheese. 



FAMILY MICROCOCCACEAE 285 

Genus III. Sarcina Goodsir.* 

(Goodsir, Edinborough Med. and Surg. Jour., 1842, 430; Lactosarcina Beijerinck, 
Arch, neerl. d. sci. exact., Ser. 2, 13, 1908, 359; Urosarcina Miquel, Ann. Microg., 1, 
1888, 517; Planosarcina Migula, Arb. Bakt. Inst. Karlsruhe, 1, 1894, 236; Pseudosar- 
cina Lohnis, Handb. d. landwirtsch. Bakt., 1910, 449 (Pseudo-sarcina, Maze, Compt. 
rend. Acad. Sci. Paris, 137, 1903, 887); Sporosarcina Orla-Jensen, Cent. f. Bakt., II 
Abt., 22, 1909, 340; Paulosarcina Enderlein, Sitzber. Ge.sell. Xaturf., Berlin. 1917, 
319 ; Phacelium Enderlein, ibid.,Sl9 ; Zymosarchia Smit, Die Garungssarcinen. Pflan- 
zenforschung. Heft 14, 1930, 26; Butyrisarcina Kluyver and Van Niel, Cent, f . Bakt., 
II Abt., 94, 1936, 400; Methanosarcina Kluyver and Van Niel, ■ibid.) From Latin 
sarcina, packet, bundle. 

Division occurs, under favorable conditions, in three planes, producing regular 
packets. Usually Gram-positive. Growth on agar abundant, usually with formation 
of yellow or orange pigment. Glucose broth slightly acid, lactose broth generally 
neutral. Gelatin frequently liquefied. Nitrites may or may not be produced from 
nitrates. Saprophytes and facultative parasites. 

The type species is Sarcina ventriculi Goodsir. 

Key to the species of genus Sarcina. 

I. Microaerophilic to anaerobic. 

A. No growth without sugars. Do not produce methane. Sub-genus Zymo- 

sarcina Smit (Die Garungssarcinen. Pflanzenforschung, Heft 14, 1930, 26). 

1. Cellulose reaction positive. Slow coagulation in litmus milk. 

1. Sarcina ventriculi. 

2. Cellulose reaction negative. Litmus milk not coagulated. 

2. Sarcina maxima. 

B. Does not utilize sugars. Produces methane. Sub-genus Methanosarcina 

Kluyver and van Niel (Cent. f. Bakt., II Abt., 94, 1936, 400). 

3. Sarcina methanica. 
II. Aerobic. 

A. No endospores present. Sub-genus Sarcinococcus subgen. nov. 

1. Not halophilic. 

a. Non-motile. 

b. Yellow pigment produced. Nitrites not produced from nitrates. 
c. Milk alkaline; coagulated. 

4. Sarcina lutea. 
cc. Milk alkaline; not coagulated. 

5. Sarcina jlava. 

bb. Orange pigment produced. Nitrites produced from nitrates. 

6. Sarcina aurantiaca. 
aa. Motile. 

7. Sarcina citrea. 

2. Halophilic red chromogen. 

8. Sarcina littoralis. 

B. Endospores present. Motile. Sub-genus Sporosarcina Orla-Jensen (Cent. f. 

Bakt., II Abt., 22, 1909, 340). 

9. Sarcina ureae. 

* Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva, 
New York, March, 1943. 



286 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



1. Sarcina ventriculi Goodsir. (Good- 
sir, Edinborough Med. and Surg. Jour., 
57, 1842, 430; Merismopedia goodsirii 
Husemann, De anim. et veget., 18 — , 
13; Merismopedia ventriculi Robin, His- 
toire des veget. parasites, 1853, 331; 
Anaerobic sarcina, Beijerinck, Proc. of 
Section of Sciences, Kon. Akad. v. We- 
tensch., Amsterdam, 7, 1905, 580; 
Zytnosarcina ventriculi Smit, Die Gar- 
ungssarcinen. Pflanzenforschung, Jena, 
Heft 14, 1930, 26; Sarcina beijerincki 
Prevot, Ann. Sci. Nat., Ser. Bot., 15, 
1933, 205.) From Latin, ventriculus, 
the stomach. 

Description taken in part from Smit 
(loc. cit.). 

Large spheres: 3.5 to 4.0 microns, 
occurring in packets of 8, 16, 32 or more 
elements. Non-motile. Gram-positive. 
Cellulose reaction positive. 

Growth occurs only in sugar media, 
containing peptones. 

Gelatin : No liquefaction. 

Deep glucose agar colonies: Alulti- 
lenticular, surrounded by a cloudy zone. 
Abundant gas. 

Glucose agar slant: Round, whitish 
colonies, several millimeters in diameter. 

Glucose broth: Abundant, flaky 
growth. Abundant gas. Acid. No 
turbidity. 

Plain peptone water: No growth. 

Sugar peptone water: Abundant 
growth. Gas. Indole not formed. 

Milk: Slow growth. Acid and co- 
agulation . 

Coagulated proteins not attacked. 

Acid and gas from glucose, fructose, 
sucrose, maltose, lactose and galactose. 
No acid from xylose, arabinose, raffinose, 
mannitol, dulcitol, salicin, starch, glyc- 
erin and inulin. 

Neutral red broth changed to fluores- 
cent yellow. 

Utilizes peptones, wort and yeast 
water as sources of nitrogen. Cannot 
utilize amino acids or inorganic nitrogen. 

Principal products of metabolism arc 
carbon dio.xide and ethyl alcohol. 



Nitrites not produced from nitrates. 

Non-pathogenic . 

Optimum pH 1.5 to 5.0. Limits of 
pH 0.9 to 9.8. 

Temperature relations : Optimum 30°C. 
Maximum 45°C. Minimum 10°C. Killed 
in ten minutes at 65°C. 

Microaerophilic to anaerobic. 

Source : Isolated from a diseased 
stomach. 

Habitat : Garden soil, dust, sand, mud; 
the stomach. 

2. Sarcina maxima Lindner. (Lind- 
ner, Die Sarcina-Organismen der Gar- 
ungsgewerbe. Inaug. Diss., Berlin, 1888, 
54; Also abstract in Cent. f. Bakt., 
4, 1888, 427; Zytnosarcina maxi'rna Smit, 
Die Giirungssarcinen. Pflanzenforsch- 
ung, Heft 14, 1930, 22; Butyrisarcina 
maxima Kluyver and van Niel, Cent. f. 
Bakt., II Abt., 94, 1936, 400.) From 
Latin maxima, largest. 

Description from Weinberg, Nativelle 
and Prevot, Les Microbes Anaerobies, 
1937, 1030 and from Smit, loc. cit. 

Large spheres: 4.0 to 4.5 microns, oc- 
curring in regular packets of 8, 16, 32 or 
more elements. Non-motile. Gram- 

positive. 

Growth occurs only in sugar media, 
containing peptones. 

Gelatin : No liquefaction. 

Deep glucose agar colonies: Multi- 
lenticular. Abundant gas produced. 

Glucose agar slant : Round, whitish 
colonies. 

Glucose broth: Abundant growth, 
flaky, gaseous, marked acidification. Dis- 
agreeable butyric odor. No turbidity. 

Sugar peptone water : Abundant 
growth, flaky, gaseous, followed by acid- 
ification. 

Milk : Not coagulated. 

Coagulated proteins not attacked. 

Cellulose reaction negative. 

Acid and gas from glucose, fructose, 
galactose, maltose, sucrose and lactose. 

Neutral red broth changed to fluores- 
cent yellow. 



FAMILY MICROCOCCACEAE 



287 



Utilizes peptones, yeast water or brotli 
as source of nitrogen. Cannot utilize 
amino acids or inorganic nitrogen. 

Principal products of metabolism are 
carbon dioxide, butyric and acetic acids. 

Non-pathogenic. 

Limits of pH 1.0 to 9.5. 

Temperature relations : Optimum 30°C. 
Maximum 40°C. Minimum 15°C. Killed 
in twenty minutes at 55°C. 

Microaerophilic to anaerobic. 

Source : Isolated from fermenting malt 
mash. 

Habitat : Acidified flour pastes, wheat 
bran; seldom in soils. Also intestinal 
contents of guinea pigs (Crecelius and 
Rettger, Jour. Bact., 46, 1943, 10). 

3. Sarcina methanica (Smit) Wein- 
berg et al. (Methaansarcine, Sohngen, 
Inaug. Diss., Delft, 1906, 104; Zymosar- 
cina methanica Smit, Die Garungssar- 
cinen. Pflanzenforschung, Heft 14, 1930, 
25; Methanosarcina methanica Kluyver 
and Van Kiel, Cent. f. Bakt., II Abt., 94, 
1936, 400; Barker, Arch. f. Mikrobiol., 
7, 1936, 420; Weinberg, Xativelle and 
Prevot, Les Microbes Anaerobies, 1937, 
1032.) From M. L. methanum, meth- 
ane; M. L. methanicus, related to 
methane. 

Description from Weinberg, Nativelle 
and Prevot {loc. cit.) and Smit {loc. cit.). 

Spheres: 2.0 to 2.5 microns, occurring 
in packets of 8 or more cocci. Non-mo- 
tile. Gram-variable. 

Growth in solutions of calcium acetate 
and possibly butyrate and inorganic am- 
monium salts. Carbon dioxide is needed 
for growth. 

In acetate-agar (with addition of some 
HaS and NaHCOs) : Colonies of 50 to 100 
microns are formed, showing gas forma- 
tion. 

Cultural characters as yet unknown. 

Peptones not attacked. 

Cellulose reaction negative. 

Utilizes ammonium salts as source of 
nitrogen. No organic nitrogen com- 
pounds utilized. 



Carbohydrates not fermented. Ethyl 
alcohol is not fermented. 

Principal products from the metab- 
olism of calcium acetate and butyrate 
are methane, carbon dioxide and calcium 
carbonate. 

Non-pathogenic . 

Optimum temperature 35° to 37°C. 

Strict anaerobe. Killed by a short 
contact with the air. 

Distinctive characters : Utilizes am- 
monium salts and acyclic acids producing 
methane and carbonic acid. 

Source : Sediment in methane fer- 
mentation (Weinberg et al.). Isolated 
from mud (Smit). 

Habitat : Swamp waters and mud ; 
fermenting sewage sludge. 

4. Sarcina lutea Schroeter. (Kryp- 
tog. Flora v. Schlesien, 8, 1, 1886, 154; 
also see Klein, Microorganisms and Dis- 
ease, 1885, 43; Eisenberg, Bakt. Diag., 
1 Aufl., Taf. 2, 1886; Fliigge, Die Mikro- 
organismen, 2 Aufl., 1886, 179; Frankland 
and Frankland, Phil. Trans. London, 
178, B, 1888, 265.) From Latin lutens, 
yellow. 

Spheres: 1.0 to 1.5 microns, showing 
packets in all media. Gram-positive. 

Gelatin colonies : Circular up to 5 mm. 
in diameter, sulfur-yellow, sinking into 
the medium. 

Gelatin stab : Slow infundibuliform 
liquefaction. 

Agar colonies : Yellow, coarsely granu- 
lar, circular, raised, moist, glistening, 
entire margin. 

Agar slant : Sulfur to chrome yellow, 
smooth, soft. 

Broth : Clear with abundant yellow 
sediment. 

Litmus milk: Coagulated, becoming 
alkaline. 

Potato : Sulfur to chrome yellow, 
raised; sometimes limited growth. 

Slight indole formation. 

Nitrites generally produced from ni- 
trates. 



288 



MANUAL or DETERMINATIVE BACTERIOLOGY 



No acid from glucose, lactose or su- 
crose. 

Hydrogen sulfide is formed. 

Aerobic. 

Optimum temperature 25°C. 

Habitat : Air. soil and water, skin 
surfaces. 

5. Sarcina flava De Bary. (Vorlesun- 
gen liber Bakterien, 1887, 151 ; Sarcina 
liquefaciens Frankland and Frankland, 
Philos. Trans. Roy. Soc. London, 178, B, 
1888, 267.) From Latin, flavus, yellow. 

Spheres: 1.0 to 2.0 microns, occurring 
in packets of 16 to 32 cells. Gram-posi- 
tive. 

Gelatin colonies: Small, circular, yel- 
lowish. 

Gelatin stab: Slowly liquefied. 

Agar slant: Yellow streak. 

Broth: Slowly becoming turbid with 
whitish, later yellowish sediment. 

Litmus milk: Alkaline, not coagulated. 

Potato: Yellow streak. 

Indole not produced. 

Nitrites not produced from nitrates. 

Aerobic. 

Optimum temperature 30° to 35°C. 

Habitat : Air, water, soil. 

6. Sarcina aurantiaca Flugge. (Die 
Mikroorganismen, 1886, 180; For descrip- 
tion see Frankland and Frankland, Phil. 
Trans. Roy. Soc. London, 178, B, 1888, 
266; Paulosarcina aurantiaca Enderlein, 
Sitzungsber. Ges. Naturf. Freunde, Ber- 
lin, 1917, 319.) From M. L., aurantiacus , 
orange-colored. 

Spheres developing packets in all 
media. Gram-positive. 

Gelatin colonies: Small, circular, dark 
yellow, entire margin, sinking into the 
medium. 

Gelatin stab: Infundibuliform lique- 
faction. 

Agar slant : Slightly raised, orange 
yellow to orange red, soft, smooth. 

Broth: Flocculent turbidity, with 
abundant sediment. 

Litmus milk: Coagulation and di- 
gestion. 



Potato : Raised, yellow -orange, glisten- 
ing to dull, granular. 
Slight indole formation. 
Nitrites not produced from nitrates. 
No HjS produced. 
Aerobic. 

Optimum temperature 30°C. 
Habitat : Air and water. 

7. Sarcina citrea (Migula) Bergey et al. 
{Micrococcus agilis citreus Menge, Cent, 
f. Bakt., 12, 1892, 52; Planococcus citreus 
Migula, Arb. bakt. Inst. Karlsruhe, 1, 
Heft 1, 1894, 236; Micrococcus agilis- 
citreus Chester, Man. of Bact., 1901, 
115; Bergey et al., Manual, 1st ed., 1923, 
74.) From M. L. citreus, lemon-yellow. 

Spheres : 0.6 to 0.8 micron, occurring 
singly, in pairs and in packets. Motile, 
possessing a single flagellum. Gram- 
positive. 

Gelatin colonies: Small, circular, yel- 
lowish, entire, becoming citron-yellow 
to orange. 

Gelatin stab: No liquefaction. 

Agar colonies: Small, yellow, convex, 
entire, smooth, glistening. 

Agar slant : Abundant, yellow, plumose, 
glistening, taking on an orange color with 
age. 

Broth: Turbid. 

Potato: Abundant, yellow growth. 

Indole not formed. 

Nitrites not produced from nitrates. 

Aerobic. 

Optimum temperature 25°C. 

Habitat: Air. 

8. Sarcina littoralis Poulsen. (Poul- 
sen, Vidensk. Meddel. naturh. Foren. i 
Copenhagen, 1879-80, 231-254; Sarcina 
morrhiiae Farlow, L'. S. Fish Commis- 
sion Report for 1878, 1880, 974; Micro- 
coccus a, H0ye, Bergens Museums 
Aarbog., No. 7, 2 Hefte, 1901, 39; Micro- 
coccus litoralis Kellerman, Cent, f . Bakt., 
II Abt., 42, 1915, 399.) From Latin, 
litus (littus) -tor is. the sea shore; -alis, 
relating to. 

The relationships of the following to 
each other and to Sarcina littoralis are 
not clear: 



FAMILY mCROCOCCACEAE 



289 



Erythroconis litoralis Oersted. (J^a- 
turh. Tidsskrift, 3, 1840-41, 555; Meris- 
mopedia litoralis Rabenhorst, Flora 
Europ. Algarum, 2, 1864-65, 57; Sarcina 
littoralis Winter in Rabenhorst, Krypto- 
gamen-Flora, 1, I Abt., 1884, 50; Pedio- 
coccus litoralis Trevisan, I generi e le 
specie delle Batteriacee, Milano, 1889, 
28; Lampropedia littoralis De Toni and 
Trevisan, in Saccardo, Sylloge Fungorum, 
8, 1889, 1049.) 

Coniothecium bertherandi Megnin. 
CRevue Mycologique, 6, 1884, 197.) Sac- 
cardo and Berlese (Atti. del R. Institute 
Veneto, Ser. VI, Vol. 3) consider C. 
bertherandi to be identical with Sarcina 
littoralis, while Zopf (Die Spaltpilze, 2 
Aufl., 1884, 73; 3 Auti., 1885, 102) 
considers C. bertherandi a stage of Beg - 
giatoa roseo-persicina. 

Description taken from Lochhead (Can. 
Jour. Res., 10, 1934, 280). 

Spheres: 1.2 to 1.6 microns occurring 
singly, in pairs, in fours, in short chains, 
and in packets, the arrangement varying 
with medium, temperature, salt concen- 
tration and age of culture. Non-motile. 
Gram stain variable, with rather more 
positive than negative cells. 

No growth in ordinary media. 

Salt gelatin : Growth slow, with no 
liquefaction. 

Starch media (20 per cent salt) : Colo- 
nies usually 1 to 3 mm, round, entire, 
convex, with a waxy appearance, brick 
red with a pale border, color appearing 
gradually. 

Starch media slants (20 per cent salt) : 
Filiform, slightly raised, entire edge. 
Coral red in color. Slight decrease in 
shade as cultures age. 

Liquid media : No growth. 

Potato : In 20 per cent salt, scanty 
growth. Slight chalky pink develop- 
ment near the top. 

Indole not formed. 

Nitrates reduced to nitrites. 

Diastatic action negative. 

Aerobic, obligate. 

Halophilic, obligate, 16-32 per cent 
salt. Optimum 20-24 per cent. 



Optimum temperature 37°C. 

Source : Isolated from seashore mud 
near Copenhagen. 

Habitat: Sea water brine, or sea salt. 
Isolated from salted hides and salted 
fish. 

The following is believed by Keller- 
man {loc. cit.) to be a variety of Sarcina 
littoralis : 

Diplococcus gadidarum Beckwith. 
(Beckwith, Cent. f. Bakt., I Abt., Orig., 
60, 1911, 351; Micrococcus litoralis gadi- 
darum Kellerman, Cent. f. Bakt., II 
Abt., 4^, 1915, 400; Pediococcus gadida- 
rum Pribram, Klassification der Schizo- 
myceten, 1933, 46.) From reddened 
salted codfish. 

9. Sarcina ureae (Beijerinck) Lohnis. 
( Planosarcina ureae Beijerinck, Cent. f. 
Bakt., II Abt., 7, 1901, 52; Lohnis, Land- 
wirtsch. bakteriol. Prakticum, 1911, 138; 
Sporosarcina ureae Kluyver and van 
Niel, Cent. f. Bakt., II Abt., 94, 1936, 
401.) From Greek, urum, urine; M. L., 
urea, urea. 

Probable synonym : Sarcina psychro- 
carterica (Rubentschick) Bergey et al. 
( Urosarcina psychrocarterica Rubent- 
schick, Cent. f. Bakt., II Abt., 64, 1925, 
168; ibid., 66, 1926, 161; ibid., 67, 1926, 
167; ibid., 68, 1926, 327; Bergey et al., 
Manual, 3rd ed., 1930, 95.) 

Spheres: 0.7 to 1.2 microns, occurring 
singly, in pairs and in packets. Atypical 
endospores present. Motile, possessing 
long peritrichous flagella. Gram- 
positive. 

Gelatin colonies: Small, circular, flat, 
tough, yellowish. 

Converts urea into ammonium car- 
bonate. 

Aerobic. 

Optimum temperature 20°C. Resists 
heating to 80°C for 10 minutes. 

Source : Isolated from urine. 

Appendix : The following names appear 
in the literature, and are listed here 
chiefly for their historical interest. 



290 



MANUAL OF DETERMINATIVE -BACTERIOLOGY 



Man}' are inadequately described, and 
probably many are synonyms. 

Micrococcus aiirantiacus Pagliani, Mag- 
giora and Fratini . (Pagliani et al . , 1887 ; 
Pediococcus aurayiliacus Trevisan, I 
generi e le specie delle Batteriacee, 
Milan, 1889, 28; Merismopedia aurantiaca 
Maggioria, Giorn. Soc. ital. d'Igiene, 11, 
1889, 355; Pediococcus maggiorae De Toni 
and Trevisan, in Saccardo, Sylloge Fun- 
gorum, 8, 1889, 1051.) From skin of the 
human foot. 

Planosarcina samesii Migula. (Eine 
bewegliche Sarcine, Sames, Cent. f. 
Bakt., II Abt., 4, 1898, 664; Migula, 
Syst. d. Bakt., 2, 1900, V and 278; Sar- 
cina agilis Matzuschita, Zeit. f . Hyg., 35, 
1900, 496.) From liquid manure. Prob- 
ably identical with Sarcina ureae 
Lohnis. 

Sarcina acidificans Migula. (Sarcina 
No. VIII, Adametz, Landwirtsch. Jahrb., 
18, 1889, 243; Migula, Syst. d. Bakt., 2, 
1900, 258.) From cheese. Winslow and 
Winslow (The Systematic Relationships 
of the Coccaceae, 1908, 235) regard this 
species as a variant of Sarcina lidea 
Schroeter. 

Sarcina agilis Saito. (Jour. Coll. 
Science Imp. Univ. Tokyo, 23, 1908, 
68; abst. in Cent. f. Bakt., II Abt., 24, 
1909,228.) From dust. 

Sarcina alba Zimmermann. (Weisse 
Sarcina, Maschek, Bakt. Untersuch. d. 
Leitmeritzer Trinkwasser, 1887, 64; Zim- 
mermann, Die Bakterien unserer Trink- 
u. Nutzwasser, Chemnitz, I Reihe, 1890, 
90.) From water. Zimmermann re- 

ported the presence of spores ; subsequent 
workers failed to observe spores, even 
when working with original cultures. 

Sarcina alba var. incana Appel. (Ber. 
d. landw. Inst. Konigsberg, Heft 5, 1900, 
89; quoted from Lohnis, Cent. f. Bakt., 
II Abt., 18, 1907, 146.) Frequently 
found in milk. Closely related to Ada- 
metz's Sarcinae Nos. VII, VIII and IX. 

Sarcina albida Gruber. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 3, 1895, 256.) 
From the stomach contents of a man with 
stomach cancer. 



Sarcina alutacea Gruber (loc. cit., 
221). From leaven. 

Sarcina aurea Mace. (Traite Pra- 
tique de Bact., 2nd ed., 1892, 371; not 
Sarcina aurea Henrici, see below.) From 
lung exudate. Possesses active oscillary 
motility, but no flagella. 

Sarcina aurea Henrici. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 1, 1894, 91; 
Sarcina aurescens Gruber, ibid., Heft 3, 
1895, 263.) From Swiss cheese. Wins- 
low and Winslow {loc. cit., 233) regard 
this species as a variant of Sarcina flava 
De Bary which has acquired certain 
fermentative powers. 

Sarcina aurescens var. mucosa Jaiser; 
quoted from Pribram, Klassifikation der 
Schizomyceten, 1933, 44. 

Sarcina bicolor Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 505.) 
From the stomach of a woodpecker (Picus 
major). Winslow and Winslow {loc. cit., 
232) regard this species as identical with 
Sarcina flava De Bary. 

Sarcina butyrica Migula. (Sarcina No. 
XI, Adametz, Landwirtsch. Jahrb., 18, 
1889, 244; Migula, Syst. d. Bakt., 2, 1900, 
240.) From cheese. Winslow and Wins- 
low {loc. cit., 233) regard this as a variant 
of Sarcina flava De Bary which has ac- 
quired certain fermentative powers. 

Sarcina Candida Lindner. (Die Sar- 
cina-Organismen der Gahrungsgewerbe, 
Inaug. Diss., Berlin, 1888, 43; Abst. in 
Cent. f. Bakt., 4, 1888, 427.) From 
water reservoir of a brewery and from 
air in the vicinity of the brewery. 

Sarcina canescens Stubenrath. (Stu- 
benrath, in Lehmann and Neumann, 
Bakt. Diag., 1 Aufl., 2, 1896, 143.) Stu- 
benrath considered this a subspecies or 
variety of his Sarcina equi from which it 
differed only by its constant gray color 
on all culture media. Winslow and 
Winslow {loc. cit., 232) regard this as 
identical with Sarcina flava De Bary. 

Sarcina carnea Gruber. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 3, 1895, 278.) 
From leaven. 

Sarcina casei Migula. (Sarcina No. 
VII, Adametz, Landwirtsch. Jahrb., 18, 



FAMILY MICROCOCCACEAE 



291 



1889, 242; Migula, Syst. d. Bakt., 2, 1900, 
239.) From cheese. Winslow and Wins- 
low {loc. cit., 233) regard this species as a 
variant of Sarcina flava De Bary which 
has acquired certain fermentative 
powers. 

Sarcina caseolytica Starlc and Scheib. 
(Jour. Dairy Sci., 19, 1936, 212.) From 
butter. 

Sarcina cervina Stubenrath. (Stuben- 
rath, in Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., 2, 1896, 146.) From the 
stomach in a case of carcinoma. 

Sarcina citrea Winslow and Winslow. 
(The Systematic Relationships of the 
Coccacceae, 1908, 234; not Sarcina citrea 
Bergey et al.. Manual, 1st ed., 1923, 74.) 
This is the name given by Winslow and 
Winslow to their Type 2, the nitrate- 
reducing group of Sarcina. 

Sarcina citrina Gruber. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 3, 1895, 269.) 
From leaven. Winslow and Winslow 
{loc. cit., 235) regard this species as 
identical with Sarcina lutea Schroeter. 

Sarcina conjunctivae Bergey et al. 
(Sarcina citrea conjunctivae Verderame, 
Cent. f. Bakt., I Abt., Orig., 59, 1911, 
384; Bergey et al.. Manual, 1st ed., 1923, 
71.) From the conjunctiva. Gram- 

negative. 

Sarcina devorans Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 502.) 
From stomach contents of a sparrow 
{Passer rnontanus). 

Sarcina equi Stubenrath. (Stuben- 
rath, in Lehmann and Neumann, Bakt. 
Diag., 1 Aufl., 2, 1896, 143.) Found 
frequently in the urine of horses. Very 
similar to Sarcina lutea according to 
Stubenrath, who names three subspecies 
or varieties: Sarcina livido-httescens, S. 
canescens and S. variabilis. Winslow 
and Winslow {loc. cit., 232) regard this 
species as identical with Sarcina flava 
De Bary. 

Sarcina erythromyxa (Overbeck) Krdl. 
(Micrococcus erythromyxa Overbeck, 
Nova Acta der Leop. -Carol, 55, No, 7, 
1891 ; Kr^l, Verzeichnisderabzugebenden 
Bak.) For a description of this species, 



see Zimmermann, Die Bakterien unserer 
Trink- und Nutzwasser, Chemnitz, II, 
1894, 70. From water. Produces a red 
pigment. 

Sarcina finientaria Lehmann and Neu- 
mann. (Eine bewegliche Sarcine, Sames, 
Cent. f. Bakt., II Abt., 4, 1898, 664; 
Lehmann and Neumann, Bakt. Diag., 2 
Aufi., 2, 1899, 146; Planosarcina samesii 
Migula, Syst. d. Bakt., 2, 1900, V and 
278; Sarcina samesii Matzuschita, Bakt. 
Diag., Jena, 1902, 300.) From liquid 
manure. Exhibits active motility with 
many long flagella. Pribram (Klassifika- 
tion der Schizomyceten, 1933, 45) re- 
gards this organism as identical with 
Sarcina ureae and Sarcina mobilis. 

Sarcina flavescens Henrici. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
91.) From Swiss cheese. Winslow and 
Winslow {loc. cit., 232) regard this species 
as identical with Sarcina flava De Bary. 

Sarcina fulva Stubenrath. (Das 
Genus Sarcina, Miinchen, 1897; see 
Lehmann and Neumann, Bakt. Diag., 2 
Aufi., 2, 1899, 143.) Isolated many 
times from stomach contents and once 
from preputial smegma. Similar to Sar- 
cina ■pulmonum. 

Sarcina fusca Gruber. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 3, 1895, 282.) 
From flour. 

Sarcina fuscescens De Bary. (Vorle- 
sungen liber Bakterien, 2 Aufl., 1887, 181 
and Botan. Centralb., 1887, 34. Re- 
duced to a synonym of Sarcina ventriculi 
Goodsir by Migula, Syst. d. Bakt., 2, 
1900, 259.) From the contents of the 
stomach. 

Sarcina gasojormans Gruber. (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 3, 1895, 
270.) From leaven. Young cultures 
produce considerable gas. 

Sarcina gigantea Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 508.) 
From stomach contents of the starling 
(Sturniis vulgaris). The diameter of a 
cell is 2.05 to 2.1 microns. Winslow and 
Winslow (loc. cit., 232) regard this species 
as identical with Sarcina flava De Bary. 

Sarcina gigantea Petter. (Proc. Kon. 



292 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Akad. Wet. Amsterdam, 34, 1931, 1417; 
Thesis, Utrecht, 1932; Compt. rend. 
Acad. Sci., Paris, 196, 1933, 300.) From 
salted herring. Halophilic. 

Sarcina hamaguchiae Saito. (Cent, 
f. Bakt., II Abt., 17, 1906, 155.) From 
soy bean mash. 

Sarcina incana Gruber (loc. cit., 248). 
From leaven. 

Sarcina incarnala Gruber. {hoc. cit., 
279; Rhodococcus incarnatus Winslow 
and Rogers, Jour. Inf. Dis., 3, 1906, 
545.) From [leaven. Produces pink pig- 
ment. 

Sarcina intermedia Gruber {Loc. cit., 
277). From leaven. Winslow and 
Winslow {loc. cit., 235) regard this 
species as identical with Sarcina lutea 
Schroeter. 

Sarcina intestinalis Zopf . (Die Spalt- 
pilze, 3 Aufl., 1885, 55.) From the in- 
testines of poultry. 

Sarcina lactea Gruber. {Loc. cit., 254; 
not Sarcina lactea Bergey et al.. Manual, 
1st ed., 1923, 73.) From leaven. 

Sarcina lactis Chester. (No. 45, Conn, 
Storrs Agr. Exp. Sta. Rept., 1894, 79; 
Chester, Man. Determ. Bact., 1901, HI.) 
From fermented milk (matzoon). Wins- 
low and Winslow {loc. cit., 232) regard 
this species as identical with Sarcina 
Jlava De Bary. 

Sarcina lactis acidi Conn, Esten and 
Stocking. (Storrs Agr. Exp. Sta. Ann. 
Rept., 1907, 125.) From milk. 

Sarcina lactis albus Conn, Esten and 
Stocking {loc, cit., 124). From milk. 

Sarcina lactis aurantiaca Conn, Esten 
and Stocking {loc. cit., 125). From 
milk. 

Sarcina lactis lutea Conn, Esten and 
Stocking {loc. cit., 124). From milk. 

Sarcina lembkei Migula. (No. 24, 
Lembke, Arch. f. Hyg., 26, 1896, 316; 
Migula, Syst. d. Bakt., 2, 1900, 241.) 
From the intestine. Winslow and Wins- 
low {loc. cit., 232) regard this species as 
identical with Sarcina jlava De Bary. 

Sarcina liquejaciens Frankland and 
Frankland. (Phil. Trans. Royal Soc. 



London, 178, B, 1888, 267.) From dust. 
Also found in cheese by Henri ci (Arb. 
bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 
95). Winslow and Winslow {loc. cit., 
232) regard this species as identical with 
Sarcina flava De Bary. 

Sarcina livida Gruber {loc. cit., 297). 
From leaven. Winslow and Winslow 
{loc. cit., 235) regard this species as 
identical with Sarcina lutea Schroeter. 

Sarcina livido-lutescens Stubenrath. 
(Stubenrath, in Lehmann and Neumann, 
Bakt. Diag., 1 Aufl., 2, 1896, US; Sarcina 
lutescens Chester, Man. Determ. Bact., 
1901, 112.) From stools in a case of enter- 
itis. Stubenrath regards this as a sub- 
species or variety of his Sarcina equi. 

Sarcina loewenbergii Mace. (Une 

sarcine pathogene, Loewenberg, Ann. 
Inst. Past., 13, 1899, 358; Mace, Traite 
Pratique de Bact., 4th ed., 1901, 464.) 
From the nasal cavity in a case of ozena. 
Probably a strongly slime-forming variety 
of Sarcina tetragena, according to Leh- 
mann and Neumann, Bakt. Diag., 5 
Aufl., ;2, 1912,206. Also see Galli-Valerio, 
Cent. f. Bakt., I Abt., Orig., 47, 1908, 177, 
for discussion. 

Sarcina luteola Gruber {loc. cit., 265). 
From leaven. Winslow and Winslow 
{loc. cit., 235) regard this species as 
identical with Sarcina lutea Schroeter. 

Sarcina marginata Gruber {loc. cit., 
268 ) . From leaven . Wi nslow and Wi ns - 
low {loc. cit., 235) regard this as identical 
with Sarcina lutea Schroeter. 

Sarcina, meliflava Gruber {loc. cit., 
272). From flour. Winslow and Wins- 
low {loc. cit., 235) consider this identical 
with Sarcina lutea Schroeter. 

Sarcina minuta De Bary. (Vorlesung- 
en fiber Bakterien, 1 Aufl., 1885; Eng. 
trans., 2nd ed., 1887, 117 and 185.) 

Sarcina mirabilis Kern. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 4, 1897, 506.) 
From intestine of the yellow-hammer 
{Ember iza citrinella) and a dove {Co- 
lumba oenas). Winslow and Winslow 
{loc. cit. ,2Z2) consider this species identi- 
cal with Sarcina flava De Bary. 



FAMILY MICROCOCCACEAE 



293 



Sarcina mobilis Maurea. (Cent. f. 
Bakt., 11, 1892, 228; Planosarcina mobilis 
Migula, Arb. bakt. Inst. Karlsruhe, 1, 
1894, 236; Micrococcus tnobilis Dyar, 
Ann. N. Y. Acad. Sci., 8, 1895, 353.) 
From old ascitic fluid. Motile, each cell 
with two short flagella. 

Sarcina mucosa Sauerbeck. (Cent, 
f. Bakt., I Abt., Orig., 50, 1909, 289.) 
From pulmonary sputum. Probably a 
slime-forming variety of Sarcina tetra- 
gena, according to Lehmann and Neu- 
mann, Bakt. Diag., 5 Aufl., 2, 1912, 206. 

Sarcina nivea Henri ci. (Arb. bakt. 
Inst. Karlsruhe, 1, Heft 1, 1894, 88.) 
From cheese. 

Sarcina olens Henrici {loc. cit., 94). 
From Camembert cheese. Winslow and 
Winslow (loc. cit., 232) regard this spe- 
cies as identical with Sarcina flava De 
Bary. 

Sarcina orangea Cahanesco. (Ann. 
Inst. Past., 15, 1901, 856.) From vagina 
of a dog. 

Sarcina paludosa Schroeter. (Schroe- 
ter, in Cohu, Kryptog. Flora v. Schlesien, 
3, I, 1886, 153.) From the waste water 
of a sugar factory. Not cultivated. 

Sarcina persicina Gruber {loc. cit., 
281). From leaven. 

Sarcina pseudogonorrhoeae Lehmann 
and Neumann. (Eine neue Sarcina, Na- 
gano, Cent. f. Bakt., I Abt., Orig., 32, 
1902, 327; Lehmann and Neumann, Bakt. 
Diag., 4 Aufl., S, 1907, 212; Sarcina naga- 
noi Mace, Traite Pratique de Bact., 6th 
Gd., 1, 1912, 631.) From pus. 

Sarcina pulchra Henrici (loc. cit., 89). 
From cheese. 

Sarcina pulmonum Hauser. (Lungen- 
sarcine, Virchow, Arch. f. path. Anat., 
9, 1856, 557; Hauser, Deutsch. Arch. f. 
klin. Medizin, 4^, 1887, 127; Sarcina 
virchowii Trevisan, Atti dell' Accad. 
Fisio-Medico-Statistica di Milano, Ser. 
IV, 3, 1885, 119.) From the sputum of a 
patient with phthisis. Motile (Leh- 
mann and Neumann, Bakt. Diag., 2 
Aufl., 2, 1899, 141). Hauser says this 
sarcina forms endogenous spores which 
may be demonstrated by Neisser's 



method of staining, and which have great 
resistance to heat. When cultivated in 
urine, it causes ammoniacal fermentation 
of the urea. Regarded by Jan Smit 
(personal communication, 1939) as identi- 
cal with Sarcina ureae. 

Sarcina radiata Kern (loc. cit., 53). 
From stomach and intestines of the rock 
dove (Columba livia) and a sparrow 
(Passer montanus). Winslow and Wins- 
low (loc. cit., 232) regard this as identical 
with Sarcina flava De Bary. 

Sarcina rosacea Migula. (Sarcina ro- 
sea (Schroeter) Lindner, Inaug. Diss., 
Berlin, 1888, 45; Migula, Syst. d. Bakt., 
2, 1900, 263.) Found frequently in dust 
and water. Lindner believed his culture 
to be Sarcina rosea Schroeter. 

Sarcina rubra Migula. (Eine rothe 
Sarcine, Menge, Cent. f. Bakt., 6, 1889, 
596; Migula, Syst. d. Bakt., 2, 1900, 
261.) From red milk. 

Sarcina schaudinni (Wolff) Pribram. 
(Planosarcina schaudinni Wolff, Cent, 
f. Bakt., II Abt., 18, 1907, 9; Pribram, 
Klassifikation der Schizomyceten, 1933, 
45.) From rotten places on potatoes. 
A motile coccus with long flagella. 

Sarcina solani Reinke and Berthold. 
(Die Zersetzung der Kartoffel durch 
Pilze, Berlin, 1879; see O. Appel in Lafar, 
Handbuch der Technischen Mykologie, 
2, 1905-08, 350.) Found in wet rotting 
of potatoes. 

Sarcina striata Gruber (loc. cit., 271). 
From flour. Winslow and Winslow (loc. 
cit., 235) regard this species as identical 
with Sarcina lutea Schroeter. 

Sarcina subflava Ravenel. (^lemoirs 
Nat. Acad. Sci., 8, 1896, 10.) From soil. 

Sarcina sulfurea Henrici (loc. cit., 
90). From cheese. Winslow and Wins- 
low (loc. cit., 235) consider this species 
identical with Sarcina lutea Schroeter. 

Sarcina superba Henrici (loc. cit., 93). 
From cheese. Winslow and Winslow 
(loc. cit., 232) regard this species as 
identical with Sarcina flava De Bary. 

Sarcina symbiotica Pribram. (Eine 
gelbe Sarcina, Gropenfiesser, Cent. f. 



294 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Bakt., II Abt., 61, 1925, 495; Pribram, 
Klassifikation der Schizomyceten, 1933, 
45.) Lives symbiotically with cock- 
roaches . 

Sarcina thermodurica Wainess and 
Parfitt. (Jour. Bact., 40, 1940, 157.) 
From milking machines and other dairy 
farm utensils. Resists pasteurization 
temperatures. 

Sarcina thermophila Bargagli-Petrucci. 
(Nuov. Giorn. Bot. Ital., 20, 1913; Abst. 
in Cent. f. Bakt., II Abt., 4S, 1915, 294.) 
From the borax-yielding waters of Tus- 
cany. Grows at temperatures up to 
75°C. 

Sarcina urinae Welcker. {Sarcina renis 
Hepworth, Microscop. Jour., 5, 1857, 1; 
Welcker, in Henle and Pfeffer, Ztschr. f. 
rat. Med., 3 Ser., 5, 1859, 199; Merismo- 
pedia vrinae Rabenhorst, Flor. europ. 
algarum, 2, 1865, 59.) Observed in the 
bladder. See below, Sarcina welcker i. 

Sarcina variabilis Stubenrath. (Stu- 
benrath, in Lehmann and Neumann, 
Bakt. Diag., 1 Autt., 2, 1896, 143.) From 
gastric contents. May be considered a 
subspecies of Sarcina equi Stubenrath. 
Winslow and Winslow {loc. cit., 232) re- 



gard this as identical with Sarcina fiava 
De Bary. 

Sarcina variegaia Pansini. (Arch. f. 
path. Anat., 122, 1890, 459.) Found 
in sputum from cases of influenza. 

Sarcina velutina Gruber {loc. cit., 
275) . From leaven. Winslow and Wins- 
low {loc. cit., 235) consider this species 
identical with Sarcina lutea Schroeter. 

Sarcina vermicularis Gruber {loc. cit., 
253). From wheat flour. 

Sarcina vermiformis Gruber {loc. cit., 
266) . From leaven. Winslow and Wins- 
low {loc. cit., 235) consider this species 
identical with Sarcina lutea Schroeter. 

Sarcina viridis flavescens Rosenthal. 
(Inaug. Diss., Erlangen, 1893; Abst. in 
Cent. f. Bakt., 16, 1894, 1024.) From 
the oral cavity. 

Sarcina welckeri Rossmann. (Ross- 
man, Ueber Urinsarcina, Flora, 40, 1857, 
641; Merismopedia welckeri Rabenhorst, 
Flora europaea, Alg. II, 1865, 58.) From 
the urinary bladder. 

Urosarcina dimorpha Beijerinck. 
(Cent. f. Bakt., II Abt., 7, 1901, 53.) 
Reported to form spores. Non-motile. 
From garden earth. 



i^AMiLY NEISSERIACEAE 295 

FAMILY VI. NEISSERIACEAE PREVOT.* 

(Ann. Sci. Nat., Ser. Bot., 15, 1933, 119.) 

Cells spherical, in pairs or in masses. Non-motile. Gram-negative. Pigment 
formation rare. The family contains aerobic and anaerobic species. Some grow poorly 
or not at all without mammalian body fluids. Optimum temperature 37°C. All 
known species are parasitic. 

Key to the genera of family Neisseriaceae. 

I. Occurring in pairs, with adjacent sides usually flattened. Aerobes, faculta- 
tive anaerobes and anaerobes. Approximately 1 micron in diameter. 

Genus I. Neisseria, p. 295. 
II. Occurring in masses, rarely in pairs. Anaerobes. Less than .5 micron in 
diameter. 

Genus II. Veillonella, p. 302. 

Genus I. Neisseria Trevisan. 

(Trevisan, Atti della Accademia Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 
105; Gonococcus Lindau (?), Just's Bot. Jahresber., I Abt., Orig., 26, 1898, 100.) 
Named for Dr. Albert Neisser who discovered the organism causing gonorrhoea in 1879. 

Paired, Gram-negative cocci with adjacent sides flattened. Four of the eleven 
species produce yellow pigment. Aerobic and anaerobic species occur. Growth on 
standard media may be poor. Biochemical activities are limited. Few carbohy- 
drates are utilized. Indole is not produced. Nitrates are not reduced. Catalase is 
produced abundantly. Parasites of mammals so far as known. 

The type species is Neisseria gonorrhoeae Trevisan. 

Key to the species of genus Neisseria. 

I. Aerobes, facultative anaerobes. 

A. Grow best on special culture media or on plain agar containing blood, blood 

serum or similar enrichment fluids, especially with added glucose. 
Grow best at 35° to 37°C; no growth below 25° or above 40°C. Not 
chromogenic. 

1. Acid from glucose, not from malto.se. Growth anaerobically. 

1. Neisseria gonorrhoeae. 

2. Acid from glucose and maltose. No growth anaerobically. 

2. Neisseria meningitidis. 

B. Grow well on ordinarj^ culture media. Grow well at 22°C. 
1. Non-chromogenic. 

a. Moist colonies on agar. No action on glucose, sucrose or mannitol. 

3. Neisseria catarrhalis. 

aa. Dry crumbly colonies on agar. Acid from glucose and sucrose; 
but not from mannitol. 

4. Neisseria sicca. 

* Revised by Prof. E. G. D. Murray, McGill University, Montreal, P.Q., Canada 
in consultation with Dr. Sara E. Branham, United States Public Health Service, 
Washington, D. C, June, 1938; further revision, August, 1943. Descriptions of 
anaerobic species reviewed by Dr. Ivan C. Hall, New York City, January, 1944. 



296 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



2. Chromogeuesis best seen on Loffler's serum, 
a. Acid from fructose, 
b. Acid from sucrose. 

5. Neisseria perflava. 
bb. No acid from sucrose. 

6. Neisseria flava. 
aa. No acid from fructose. 

b. Acid from glucose. 

7. Neisseria subflava. 
bb. No acid from glucose. 

8. Neisseria flavescens. 
II. Anaerobes. 

A. Gas produced from peptone broth. 

9. Neisseria discoides. 

B. No gas produced. 

1. Odor of rancid butter. 

10. Neisseria reniformis . 

2. No rancid odor. 

11. Neisseria orbiculata. 



1. Neisseria gonorrhoeae Trevisan. 
(Micrococcus der Gonorrhoe, Neisser, 
Vorl. Mitteil., Cent. f. Medicinische 
Wissenschaft, 17, 1879, 497; Trevisan, 
Atti della Accademia Fisio-Medico-Sta- 
tistica in Milano, Ser. 4, 3, 1885, 105.) 
From Greelc, gonorrhoea, flux of semen; 
M.L. genitive of gonorrlioea. 

Synonyms: Gonococcus, Diplococcus 
der Gonorrhoe, Bumm, Der Mikroorgan- 
ismen der gonorrhoisclien Sclileimhauter- 
krankung, Weisbaden, 1885, 16; Merismo- 
pedia gonorrhoeae Zopf, Die Spaltpilze, 
1885, 54 ; Micrococcus gonorrhoeae Fliigge, 
Die Mikroorganismen, 1886, 156; Micro- 
coccus gonococcus Schroeter, in Cohn, 
Kryptog. Flora v. Schlesien, S, I, 1886, 
147 ; Diplococcus gonorrhoeae Lehmann 
and Neumann, Bakt. Diag., 1 Aufl., 2, 
1896, 150; Micrococcus gonorrhoeae Leh- 
mann and Neumann, ibid., 4 Aufl., 2, 
1907, 212. 

Spheres: 0.6 to 1.0 micron, occurring 
singly and in pairs, the sides flattened 
where they are in contact. Gram- 
negative. 

Grows only on media with the addition 
of body fluids (blood, ascites, etc.), or 
other specially prepared media. 

Colonies are small, transparent, even- 



tually (2 to 4 days) developing a lobate 
margin, grayish-white with a pearly opal- 
escence by transmitted light. Larger 
colonies on special media. 

Acid from glucose. No acid from 
maltose, fructose, sucrose and mannitol. 

Optimum temperature 37°C. No 
growth below 25° or above 40°C. 

Aerobic to facultative anaerobic. 
Many strains develop more readily with 
increased CO2 tension. 

Common name : Gonococcus. 

Source: Originally found in purulent 
venereal discharges. Also found in 
blood, conjunctiva, joints and cerebro- 
spinal fluid. 

Habitat: The cause of gonorrhoea and 
other infections of man. Not found in 
other animals. 

2. Neisseria meningitidis (Albrecht 
and Ghon) Holland. (Diplokokkus intra- 
cellularis meningitidis Weichselbaum. 
Fortschr. d. Med., 5, 1887, 583 ; Neisseria 
weichselbaumii Trevisan, I generi e le 
specie delle Batteriacee, 1889, 32; not 
Diplococcus intracellularis Jaeger , Ztschr. 
f. Hyg., 19, 1895, 353; not Tetracoccus 
intracellularis Jaeger, ibid., 368; not 
Streptococcus intracellularis Lehmann 



FAMILY NEISSERIACEAE 



297 



and Neumann, Bakt. Diag., 1 Aufl., 2, 
1896, 132; Micrococcus intraccllularis 
Migula, Syst. d Bakt., 2, 1900, 189; 
Micrococcus meningococcus cerebrospi- 
nalis Albrecht and Ghon, Wiener klin. 
Wochnschr., U, 1901, 988; not Strepto- 
coccus weichselbaurnii Chester, Man. 
Determ. Bact., 1901, 64; not Meningo- 
coccus intracellularis Jaeger, Cent. f. 
Bakt., I Abt., Orig., 33, 1903, 23; Micro- 
coccus meningitidis Albrecht and Ghon, 
Cent. f. Bakt., I Abt., Orig., 33, 1903, 
498 ; Diplococcus intracellularis Weichsel- 
baum, Cent. f. Bakt., I Abt., Orig., 33, 
1903, 511; Micrococcus intracellularis 
meningitidis de Bettencourt and Franga, 
Ztschr. f . Hyg., 46, 1904, 464 ; Diplococcus 
meningitidis, ibid., 495; Holland, Jour. 
Bact., 5, 1920, 224; Neisseria intracel- 
lularis-meningitidis Holland, ibid., 224; 
Neisseria intracellularis Holland, ibid., 
224; see also Elser and Huntoon, Jour. 
Med. Res., 20 fN. S. 15), 1909, 371 and 
Murray, Med. Res. Council, London, 
Special Report Series No. 124, 1929 for 
detailed studies of the group.) From 
Greek, meninx, meninges, a membrane, a 
membrane covering the brain; M. L. 
genitive of meningitis, an inflammation 
of the meninges. 

The binomial, Neisseria intracellularis, 
used in previous editions of the Manual 
has proved confusing because the names 
Micrococcus intracellularis, Diplococcus 
intracellularis and Streptococcus intra- 
cellularis, have been used loosely foi- 
unrelated organisms . Neisseria weichsel - 
baumii has been so rarely and loosel.y 
used that any attempt to introduce it 
now is inadvisable despite rights of 
prioritJ^ The equally available name, 
Neisseria meningitidis, has therefore 
been adopted to avoid further confusion. 
It has the obvious advantage of associ- 
ation with the common name, meningo- 
coccus, which has been so freciuently used 
in the literature. 

In 1898, Councilman, Mallory antl 
Wright (Epidemic Cerebrospinal Menin- 
gitis and its Relation to Other Forms of 



Meningitis, Boston, 1898) definitely 
established the Gram-negative coccus as 
the cause of epidemic meningitis and 
clarified the confusion created because 
Jaeger regarded the coccus that he 
isolated (see Diplococcus crassus von 
Lingelsheim) as identical with Neisseria 
meningitidis. 

Spheres : 0.6 to 0.8 micron in diameter, 
occasionally larger, occurring singh-, in 
pairs with adjacent sides flattened, or 
occasionallj- in tetrads. Gram-negative. 

Good growth is obtained on media con- 
taining blood, blood serum and other 
enrichment fluids with added glucose. 
Best growth on special media. 

Blood agar plates are generally em- 
ployed to isolate the organism. The 
colonies are small, slightly convex, trans- 
parent, glistening. Colonies large on 
special media. 

Older cultures may show growth on 
neutral agar or glucose agar, properly 
prepared. Frequent transplantation is 
necessary to keep the organism alive in 
recently i.solated strains ; older strains 
survive for one month or longer at 37°C 
and for years on special media. 

Acid from glucose and maltose. No 
acid from fructose, sucrose and mannitol. 

Nitrites not produced from nitrates 
(Branham). 

Optimum temperature 37°C. No 
growth at 22° or at 40°C. 

Aerobic, no growth anaerobically. 

Common name: Meningococcus. 

Source: Originalh' found in cerebi-o- 
spinal fluid. Also found in nasopharynx , 
blood, conjunctiva, pus from joints, 
petechiae in skin, etc. 

Habitat: Nasopharynx of man, not 
found in other animals. Cause of epi- 
demic cerebrospinal fever (meningitis). 

Four main varieties or types of Neis- 
seria meningitidis have been differen- 
tiated by Gordon and Murray (Jour. 
Roy. Army :\Ied. Corps, 25 (2), 1915, 
423) and by others on the basis of agglu- 
tination reactions with immune serums. 



298 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



3. Neisseria catarrhalis (Frosch and 
Kolle) Holland. {Micrococcus catarrh- 
alis Frosch and Kolle, in Fliigge, Die 
Mikroorganismen, 3 Aufl., 2, 1896, 154; 
Diplococcus pharyngis communis von 
Lingelsheim, Klin. Jahrb., 15, 1906, 408; 
Holland, Jour. Bact., 5, 1920, 224.) 
From Greek, catarrh, a running down. 

Spheres : 0.6 to 0.8 micron in diameter, 
occurring singly or in pairs with adjacent 
sides flattened, occasionally in fours. 
Gram-negative. 

Agar colonies: Small, circular, grayish 
white to dirty white, with erose margins. 

Broth: Turbid, often with slight pel- 
licle. 

No acid from any of the carbohydrates. 

Optimum tempei'ature 37°C. Grows 
well at 22°C. 

Aerobic, facultative. 

Source: Nasopharynx, saliva and res- 
piratory tract. 

Habitat : Human mucous membrane of 
the respiratory tract. Often associated 
with other organisms in inflammations of 
the mucous membrane. 

Note: Topley and Wilson (Prin. of 
Bact., 1931, 349) state that Neisseria 
pharyngis cinerea (Micrococcus pharyngis 
cinereus von Lingelsheim, Klin. Jahrb., 
15, 1906, 373) resembles Neisseria catar- 
rhalis so closely that it should probably be 
regarded as a variety of this species. 

4. Neisseria sicca (von Lingelsheim) 
Bergey et al. {Diplococcus pharyngis 
siccus von Lingelsheim, Klin. Jahrb., 15, 
1906, 409; Diplococcus siccus von Lingels- 
heim, Ztschr. f. Hyg., 59, 1908, 476; 
Micrococcus pharyngis siccus Kutscher, 
in Kolle and Wassermann, Handb. d. 
Path. Mikroorganismen, 2 Aufl., 4, 1912. 
603; Micrococcus pharrjngis-siccus Hol- 
land, Jour, of Bact., 5, 1920, 224; Neis- 
seria pharyngis-sicci (sic) Holland, ibid.; 
Bergey et al., Manual, l.st ed., 1923, 43.) 
From Latin, sicca, dry. 

Spheres : 0.6 to 0.8 micron in diameter, 
occurring singly and in pairs with ad- 
jacent sides flattened. Gram-negative. 



Blood agar colonies : Grayish, somewhat 
dry, crumbling when an effort is made to 
remove them. 

Ascitic agar colonies: Small, very firm 
and adherent to medium, becoming cor- 
rugated on the surface. 

The organisms precipitate spontaneous- 
ly when suspended in normal salt solu- 
tion. 

Acid from glucose, fructose, maltose 
and sucrose. No acid from mannitol. 

Optimum temperature 37°C. Grows 
at 22°C. 

Aerobic, facultative. 

Source: Nasopharynx, saliva and spu- 
tum. 

Habitat : Mucous membrane of the 
respiratory tract of man. 

5. Neisseria perflava Bergey et al. 
(Chromogenic group I, Elser and Hun- 
toon, Jour. Med. Res., .20 (N. S. 15), 1909, 
415; Bergey et al.. Manual, 1st ed., 1923, 
43.) From Latin per, very and flavus, 
yellow. 

Spheres: 0.6 to 0.8 micron, occurring 
singly and in pairs with adjacent sides 
flattened. Gram-negative. 

Glucose agar colonies: Small, circular, 
slightly raised, greenish-gray by reflected 
light, and greenish-yellow and semi- 
opaque by transmitted light. The sur- 
face is smooth, glistening. The growth 
is adherent to the medium. Chromo- 
genesis best seen on Loffler's blood serum 
medium. 

Ascitic agar colonies : Like those on 
glucose agar. 

Acid from glucose, maltose, fructose, 
sucrose and mannitol. 

Optimum temperature 37°C. Grows 
at22°C. 

Aerobic, facultative. 

Source : Nasopharynx, saliva and spu- 
tum. 

Habitat : Mucous membrane of respira- 
tory tract of man. 

6. Neisseria flava Bergey etal. {Dip- 
lococcus pharyngis flaims I and possibly 



FAMILY NEISSERIACEAE 



299 



Diplococcus pharyngis flaws II, von 
Lingelsheim, Klin. Jahrb., 15, 1906, 409; 
Diplococcus flavus /and possibly Diplo- 
coccus flavus II, V. Lingelsheim, Zeitschr. 
f. Hyg., 59, 1908, 476; Micrococcus 
pharyngis flavus I and possibly Micro- 
coccus pharyngis flavus II, Lehmann and 
Neumann, Bakt. Diag., 7 Aufi., 2, 1927, 
259; Chromogenic group II, Elser and 
Huntoon, Jour. Med. Res.,^0 (N. S. 15), 
1909, 415; Bergey et al.. Manual, 1st ed., 
1923, 43.) From Latin flavus, yellow. 

Spheres: 0.5 to 0.8 micron, occurring 
singly and in pairs with adjacent sides 
flattened. Gram-negative. 

Glucose agar colonies: Small, circular, 
slightly raised, greenish-gray by reflected 
light and greenish-yellow by transmitted 
light. Growth not adherent to medium. 
Surface colony is smooth with numerous, 
rather coarse crumbs in center. Margin 
entire, or rarely slightlj" irregular. 
Chromogenesis best seen on Loffier's 
blood serum medium. 

Ascitic agar colonies : Like those on 
glucose agar. 

Acid from glucose, fructose and 
maltose. No acid from sucrose or man- 
nitol. 

Optimum temperature 37°C. Grows 
at22°C. 

Source: Nasopharynx, cerebro-spinal 
fluid in cases of meningitis (very rare). 

Habitat : Mucous membrane of respira- 
tory tract. 

7. Neisseria subfiava Bergey et al. 
(Chromogenic group III, Elser and 
Huntoon, Jour. Med. Research, 20 (N.S. 
15), 1909, 415; Bergey et al.. Manual, 1st 
ed., 1923, 44.) From Latin sub, less and 
flavus, yellow. 

Spheres: 0.6 to 0.8 micron, occurring 
singly and in pairs with adjacent sides 
flattened. Gram-negative. 

Glucose agar colonies: Small, slightly 
raised, pale greenish-yellow, especially 
on primary culture. 

Acid from glucose and maltose. No 
acid from fructose, sucrose or mannitol. 



Agglutinates in normal rabbit serum. 

Optimum temperature 37°C. Little 
or no growth at 22°C. 

Aerobic, facultative. 

Easily confused with Xeisseria men- 
ingitidis. 

Source: Nasopharynx. 

Habitat : Mucous membrane of the 
respiratory tract of man. 

8. Neisseria flavescens Branham. 
(U. S. Public Health Service, Pub. 
Health Repts., 43, 1930, 845.) From 
Latin flavescens, becoming yellow. 

Biscuit-shaped cocci occurring in flat- 
tened pairs. Giant forms common. 
Gram -negative. 

Glucose agar : Poor growth. 

Blood agar : Good growth, colonies less 
moist than those of the meningococcus. 
Golden yellow pigment. Greenish-yel- 
low on Loffler's blood serum medium. 

Semisolid agar: Good growth. 

No acid from anj' of the carbohydrates. 

Optimum temperature 37°C. 

Aerobic, facultative. 

Serologically homogeneous group. 

Source : Cerebro-spinal fluid in cases 
of meningitis. 

Habitat : Probably mucous membrane 
of respiratory tract of man. 

Note : Wilson and Smith (Jour. Path, 
and Bact., 31, 1928, 597) do not regard 
differences in sugar fermentations, 
chromogenesis, appearance of colonies, 
etc. sufficiently constant to warrant the 
separation of the species Neisseria catar- 
rhalis, N. flava, N. cinerea, N. mucosa 
and N. sicca. They recommend that all 
be grouped under a single species known 
as Neisseria pharyngis (Diplococcus 
pharyngis) . 

9. Neisseria discoides Prevot. (Ann. 
Sci. Nat., Ser. Bot., 15, 1933, 106.) 
From Greek, discoeides, discus shaped; 
Latin adj., disk-shaped. 

Spheres: 0.6 to 0.7 micron, occurring 
in pairs or tetrads. Gram-negative. 



300 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Gelatin: No liquefactiou. 

Deep agar colonies : Lenticular, up to 
1 mm in diameter. Grows in a narrow 
disk about 1 cm below the surface. Gas 
produced. 

Broth : Turbid. Fine granular precipi- 
tate. Slight rancid odor and inflam- 
mable, explosive gas produced. 

Peptone water: Gas produced. 

Indole not formed. 

No action on milk. 

Coagulated proteins not digested. 

Carbohydrates not attacked. 

Hydrogen sulfide not produced. 

Neutral red glucose broth: Becomes 
pink, but no further change. 

Optimum pH 7.0 to 8.0. 

Temperature relations : Optimum 37°C. 
No growth at 28°C. Killed in half an 
hour at 60°C. 

Non-pathogenic. 

Strict anaerobe. 

Distinctive characters: Colonies grow 
in narrow zone 1 cm below the surface 
of an agar stab; gas produced from pep- 
tones. 

Source : Isolated from bronchial mucus, 
respiratory system ; dental and tonsillary 
focal infections. 

Habitat: Buccal cavity (human) and 
probably also in other warm-blooded 
animals. 

10. Neisseria reniformis (Cottet) Tre- 
vot. {Diplococcus reniformis Cottet, 
Compt. rend. Soc. Biol., 52, 1900, 421; 
Micrococcus reniformis Oliver and Wher- 
ry, Jour. Inf. Dis., 28, 1921, 341 ; Prevot, 
Ann. Sci. Nat., Ser. Bot., 15, 1933, 102.) 
From Latin, ren (renes) , kidney ; -formis , 
form, i.e. kidney-shaped. 

Spheres :0.8 to 1 .0 micron, bean-shaped, 
occurring in pairs. Gram-negative. 

Gelatin : No liquefaction. 

Deep agar colonies : Appear in 24 to 48 
hours; at first punctiform, then lenti- 
cular; small, 0.3 to 0.5 mm. No gas 
produced. 

Agar slant : Minute, bluish-white, dew- 
drop colonies. 



Broth : Turbid in 24 hours ; flocculent 
precipitate rapidly formed, clearing the 
medium. No gas produced, but a rancid 
odor is present. 

Peptone water: Very meagre growth. 
Traces of indole. 

Milk: Unchanged. 

Coagulated proteins not digested. 

Slight amount of acid from glucose by 
one strain only. 

Optimum pH 7.0. Limits of pH 6.0 
to 8.0. 

Temperature relations : Optimum 37°C. 
No growth at 22°C. Killed in half an 
hour at 60°C. or in an hour at 56°C. 

Pathogenic. 

Strict anaerobe. 

Distinctive character : Odor of rancid 
butter. 

Source : Isolated in several cases from 
suppurations of the urino-genital system. 

Habitat : Presumably in bodies of 
warm-blooded animals. 

11. Neisseria orbiculata Prevot. (Dip- 
lococcus orbicuhts Tissier, Ann. Inst. 
Past., 22, 1908, 204; Prevot, Ann. Sci. 
Nat., Ser. Bot., 15, 1933, 109.) From 
Latin, orbiculatus, having the form of 
an orb or sphere. 

Spheres: 1.5 to 2.0 microns, occurring 
in pairs. Gram-negative. 

Gelatin: No growth at 22°C. 

Deep agar colonies : After 36 to 48 hours , 
large, lenticular, very regular, whitish, 
almost transparent. Gas not produced. 

Broth : Turbid. Sediment. 

Milk : No coagulation. 

Egg white not attacked. 

Proteoses attacked without formation 
of indole. 

Acid from glucose. Acid produced 
feebly from lactose. No acid from su- 
crose. 

Temperature relations : Optimum 37°C. 
No growth at 22°C. Killed at 60°C. 

Non-pathogenic. 

Strict anaerobe. 

Distinctive characters : Large size ; no 
gas production . 



FAMILY NEISSERIACEAE 



301 



Source : Isolated from feces of young 
children. 

Habitat: Intestinal canal. Not com- 
mon. 

Appendix I: Additional species have 
been placed in this genus as given below. 
Some are undoubtedly identical with 
previously described species, while some 
may belong in other genera. 

Diplococcus crassus von Lingelsheim. 
{Diplococcus intracellular is Jaeger, 
Ztsch. f. Hyg., 19, 1893, 353; Tetracoccus 
intracellular is, ibid., 318; von Lingels- 
heim, Ztschr. f. Hyg., 59, 1908, 467; 
Micrococcus crassus Lehmann and Neu- 
mann, Bakt. Diag., 7 Aufl., 2, 1927, 
259.) Commonly found in nasopharyn- 
geal secretions, also in the cerebrospinal 
fluid of suspected cases of meningitis. 
Also known as Jaeger's coccus or as 
Jaegersher Modifikation der Meningo- 
coccus. 

Diplococcris mucosus von Lingelsheim. 
(von Lingelsheim, Klin. Jahrb., 15, 1906, 
373, and Ztschr. f. Hyg., 59, 1908, 457; 
Neisseria mucosa Murray, in Manual, 5th 
ed., 1939, 283; not Streptococcus mucosus 
Howard and Perkins, Jour. Med. Res., 
6, (N.S. 1), 1901, 174; not Pneumococcus 
mucosus Park and Williams, Jour. Exp. 
Med., 7, 1905, 411.) From nasal secre- 
tions. This Gram-negative coccus is said 
to show similarity to the meningococcus 
and to be like the diplococcus found bj' 
Weichselbaum and Ghon (Weiner Klin. 
Wchnschr., No. 24, 1905) in nasal secre- 
tions of a healthy person. Clearly it is 
different from the Gram-positive, mu- 
coid type of pneumococcus which is de- 
scribed by Binaghi (Cent. f. Bakt., I 
Abt., 22, 1897, 273), Howard and Perkins 
(Jour. Med. Res., 6, 1901, 174), Park and 
Williams (Jour. Exp. Med., 7, 1905, 411) 
and others. 

Micrococcus pharyngis cinereus von 
Lingelsheim. (Klin. Jahrb., 15, 1906, 
373 ; Micrococcus cinereus v. Lingelsheim, 
Ztschr. f. Hyg., 59, 1908, 456; Neisseria 
cinerea Murray, in Manual, 5th ed., 1939, 



283.) From mucous membrane of nose 
and throat. 

Neisseria arthritica (Costa) Hauduroy 
et al. {Micrococcus arthritica Costa, 
Comp. rend. Soc. Biol., Paris, 85, 1920, 
933; Hauduroy, Ehringer, Urbain, Guil- 
lot and Magrou, Dictionnaire des Bac- 
teries Pathogenes, Paris, 1937, 296.) 
Isolated from a case of human arthritis. 

Neisseria edigtoni (sic) Trevisan. 
{Diplococcus scarlatinae sanguinis Jamie - 
son and Edington, Brit. Med. Jour., 1, 
1887, 1265; Trevisan, I generi e le specie 
delle Batteriacee, 1889, 32.) From a 
scarlet fever patient. 

Neisseria fulva De Bord, (Jour. Bact., 
38, 1939, 119; Iowa State Coll. Jour. Sci., 
16, 1942, 471.) From conjunctivitis 
and vaginitis. 

Neisseria gibbonsi Hauduroy et al. 
(Gram -negative coccus, Gibbons, Jour. 
Inf. Dis., 43, 1929, 289; Hauduroy et al., 
Diet. d. Bact. Path., 1937, 300.) Iso- 
lated from skin abscesses in rabbits and 
guinea pigs. 

Neisseria gigantea De Bord. (Jour. 
Bact., 38, 1939, 119; Iowa State Coll. 
Jour. Sci., 16, 1942, 472.) From a normal 
vagina. 

Neisseria luciliarum Brown. (Amer. 
Mus. Novit., No. 251, 1927, 3.) A motile. 
Gram-negative diplococcus that prob- 
ably should be placed in the genus Mi- 
crococcus. From a dead fly, Lucilia 
sericata killed by Bacillus lutzae. 

Neisseria pseudocalarrhalis Huntoon. 
(Jour. Bact., 27, 1934, 108.) Like A^ezs- 
seria catarrhalis, shows no action on 
carbohydrates but is culturally more like 
Neisseria meningitidis and forms homo- 
geneous suspensions in a salt solution. 
From nasopharynx. 

Neisseria rebellis Trevisan. (Micro- 
coccus in Trachoma folliculare, Kuchark- 
sky, 1887; Trevisan, I generi e le specie 
delle Batteriacee, Milan, 1889, 32.) 
From trachoma. 

Neisseria venezuelensis Hauduroy et 
al. (Riguez, Gaceta Med. de Caracas, 



302 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



June 30, 1935; Pedro del Corral, Rev. de 
Med. y Cir. de la Clinica Macacay, April, 
1935; Hauduroy et al., Diet. d. Bact. 



Path., 1937, 308.) Found in localized 
epidemics of cerebrospinal meningitis 
in Venezuela. 



, Genus II. Veillonella Prevot.* 

(Ann. Sci. Nat., Ser. Bot., 15, 1933, 118.) Named for A. Veillon, the French 
bacteriologist, who isolated the type species. 

Small, Gram-negative cocci averaging 0.3 micron. Occur in masses, rarely in pairs 
or short chains. Cells undifferentiated. United by an interstitial substance of 
ectoplasmic nature. The known species are anaerobic. Good growth on standard 
culture media. Biochemical activity pronounced. Harmless parasites in mouth and 
intestine of man and animals. 

The type species is Veillonella parvnla (Veillon and Zuber) Prevot. 

Key to the species of genus Veillonella. 

I. Acid and gas from glucose. Weakly hemolytic. 

1. Veillonella parvula. 

II. Carbohydrates not attacked. Gas produced from peptone broth. Non- 
hemolytic. 

2. Veillonella gazogenes. 



1. Veillonella parvula (Veillon and 
Zuber) Prevot. (Staphylococcus parvu- 
lus Veillon and Zuber, Arch. med. Exp., 
1898, 542; Micrococcus parvulus Bergey 
et al.. Manual, 3rd ed., 1930, 92; Prevot, 
Ann. Sci. Nat., Ser. Bot., 15, 1933, 119.) 
From Latin, parvulus, very small. 

Description from Prevot (loc. cit.). 

Very small spheres: 0.2 to 0.4 micron, 
occurring in masses, occasionally in very 
short chains. Gram-negative. 

Gelatin : No liquefaction. 

Semisolid agar (Veillon) colonies: At 
first punctiform, becoming lenticular, 
reaching a diameter of 2 mm. Gas bub- 
bles. 

Blood agar colonies : Usually surround- 
ed by a clear halo; weakly hemolytic. 

Agar slant: Transparent, bluish, min- 
ute colonies. 

Peptone broth: Turbid with fine sedi- 
ment. 

Glucose broth: Turbid. Faintly fetid 
odor. Gas produced contains CO2, H2 
and H2S. 

Broth serum: Very abundant, rapid 
growth . 



Milk: No acid. Not coagulated. 
Some strains produce gas. 

Small amount of indole formed. 

Nitrites produced from nitrates. 

.\cid and gas from glucose. Slight 
amount of acid from fructose, galactose 
and sucrose. Some strains feebly attack 
mannitol, maltose and inulin. 

Coagulated protein not attacked. 

.Ammonia not produced. 

Hydrogen sulfide produced. 

Optimum pH G.5 to 8.0. 

Temperature relations : Optimum 37°C. 
Grows feebly at 22°C. Killed in one 
hour at 55°C. 

Strict anaerobe. 

Distinctive characters : Fermentation 
of polypeptids to produce hydrogen, 
carbon dioxide, hydrogen sulfide and 
indole ; fermentation of sugars ; hemoly- 
sis of blood; production of nitrites from 
nitrates. 

Source : Isolated by Veillon and Zuber 
from appendices, buccal cavities and 
lungs. Of the 13 strains studied by 
Prevot, 3 were isolated from pulmonary 
gangrene, one from an appendix, one 



* Revised by Prof. E. G. D. Murray, McGill University, Montreal, P. Q., Can- 
ada, June, 1938. Descriptions reviewed by Dr. Ivan C. Hall, New York City, Jan- 
uary, 1944. 



FAMILY XEISSERIACEAE 



303 



from alveolar pyorrhea, 5 from amniotic 
fluid, 2 from abscesses and pulmonary 
congestion and one from the buccal 
cavity of a normal rabbit. Found in 
suppurative lesions or pus. It may oc- 
casionall}' be pathogenic and invade the 
tissues, causing suppurations, alone or 
in association with other pyogenic or- 
ganisms. 

Habitat : Normally a harmless parasite 
found in natural cavities of man and 
animals, especially the mouth and digest- 
ive tract. 

la. Veillo7ieila parvula var. minima 
Prevot. {Staphylococcus minimus Gio- 
clli, Boll. R. Accad. Med. di Genova, 
1907; Abst. in Cent. f. Bakt., I Abt., 
Ilef., 4-'-, 1908-09, 595; Micrococcus mini- 
mus Bergey et al.. Manual, 1st ed., 1923, 
69; Prevot, Ann. Sci. Nat., Ser. Bot., 
15, 1933, 125.) From Latin, minimus, 
smallest. 

Differs from VeillonelUi parvula only 
in its slightly smaller size (0.2 to 0.3 mi- 
cron). Growth only at 37°C. Nogrowth 
ongelatin. Growthonthe wall of the cul- 
ture tube in fine flakes, not clouding the 
medium, and no plasmolysis in a 5 per 
cent salt solution. 

Source : Isolated from a periuterine 
abscess. 

lb. Veillonella parvula var. branhamii 
Prevot. (Anaerobic micrococcus, Bran- 
ham, Jour. Inf. Dis., 41, 1927, 203; ibid., 
42, 1928, 230; Micrococcus branhamii 
Bergey et al.. Manual, 3rd ed., 1930, 92; 
Prevot, Ann. Sci. Nat., Ser. Bot., 15, 
1933, 126.) Named for Dr. Sara E. 
Branham, of the National Institute of 
Health, Washington, D. C. 

Serologically distinct from Veillonella 
parvula. One strain liquefied gelatin 
slowly. 

Source : Isolated from nasal washings 
in two cases of infiuenza. 

Ic. Veillonella parvula var. thomsonii 
Prevot. (Anaerobic diplococcus, Thom- 
son, Jour. Trop. Med. and Hyg., 26, 1923, 



227 and Ann. Pickett-Thomson Res. 
Lab., ;, 1924-25, 105 and 164; Prevot, 
Ann. Sci. Nat., Ser. Bot., 15, 1933, 126; 
Micrococcus thomsoni Hauduroy et al., 
Diet. d. Bact. Path., 1937, 283.) Named 
for Dr. David Thomson of London, 
England. 

Differs but slightly from Veillonella 
parvula in that it requires some accessory 
factor of growth found in serum or similar 
body fluids, testicular agar and the like. 

Source : Found in the throat in measles 
and scarlet fever. 

2. Veillonella gazogenes (Hall and 
Howitt) Murray. {Micrococcus gazo- 
genes alcalescens anaerobius Lewkowicz, 
Arch. Med. Exp., 18, 1901, 633; Micro- 
coccus gazogenes Hall and Howitt, 
Jour. Inf. Dis., 37, 1925, 112; not Micro- 
coccus gazogenes Choukevitch, Ann. 
Inst. Pasteur, 25, 1911, 356; Veillonella 
alcalescens Prevot, Ann. Sci. Nat., S6r. 
Bot., 15, 1933, 127; Micrococcus alca- 
lescens Hauduroy et al.. Diet. d. Bact. 
Path., 1937, 274; Murray, in Manual, 
5th ed., 1939, 287.) From Latin, the gas- 
producing Veillonella. 

The species name gazogenes as given by 
Hall and Howitt is well established in the 
literature for this organism. It is valid 
under the rules when the organism is 
placed in a new genus {Veillonella) in 
spite of the earlier use of Micrococcus 
gazogenes by Choukevitch for a different 
organism. 

Spheres : 0.3 to 0.7 micron, average 0.4 
micron, occurring in irregular masses, 
rarely in pairs, short chains or singly. 
Gram-negative. 

Gelatin: No liquefaction. 

Deep agar colonies : At first punctiform , 
becoming lenticular. Gas bubbles ap- 
pear after 16 to 18 hours. 

Blood agar plate : Minute colonies. 
Non-hemolytic. Several strains show 
greenish colonies. 

Peptone broth: Gas produced. Broth 
becomes slightly alkaline. 

Indole not formed. 



304 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Milk: Gas, but no acid. No co- 
agulation. 

Ammonia and hydrogen produced in 
small amounts. 

Egg-white and coagulated serum not 
attacked. 

Hydrogen sulfide not produced. 

Carbohydrates not attacked. 

Nitrites not produced from nitrates. 

Slowly plasmolysed in 5 per cent NaCl 
solution. 

Optimum pH 6.0 to 8.0. Will grow in 
broth of pH 5.5. 

Temperature relations : Optimum 37°C. 
Some strains grow at 22°C. Killed at 
56°C in one hour, or at 65°C in a half 
Qour, or at 80°C in 10 minutes. 

Non-pathogenic (Lewkowicz's strains). 
Two strains (Pr6vot) pathogenic for 
rabbits. 

Strict anaerobe. 

Distinctive characters: Differs from 
Veillonella parvula in that it does not 
ferment sugars, does not produce H2S 
nor indole, is not hemolytic, does not 
Droduce nitrites from nitrates, and does 
not develop fetid odors. 

Source: Isolated (Lewkowicz) from 
mouth of a healthy infant. Twenty -four 
strains (Hall and Howitt) from human 
saliva. Fifteen strains (Prevot) one 
from alveolar pyorrhea, one from pul- 
monary gangrene, 5 from tonsils, one from 
appendix, 2 from measles, 3 from scarlet 
fever, and 2 from normal guinea pigs and 
rabbits. 

Habitat : Prevalent in saliva of man 
and animals. 

2a. Veillonella gazogenes var. gingivalis 
Murray. (Kleiner Micrococcus, Ozaki, 
Cent. f. Bakt., I Abt., Orig., 62, 1912, 
83; Micrococcus gingivalis Bergey et al.. 
Manual, 1st ed., 1923, 69; Veillonella 
alcalescens var. gingivalis Pr6vot, Ann. 
Sci. Nat., Ser. Bot., 15, 1933, 133 ; Murray 
in Manual, 5th ed., 1939, 288.) From 
Latin, pertaining to the gums. 



Differs from Veillonella gazogenes by its 
ability to grow at 22°C, and by the fact 
that glucose favors its growth although 
this carbohydrate is not fermented. 

Source: Oral cavity and (Prevot) two 
strains from the intestine. 

2b. Veillo7iella gazogenes var. minutis- 
sima Murray. (Micrococcus minutissi- 
mus Oliver and Wherry, Jour. Inf. Dis., 
28, 1921, 342; Veillonella alcalescens var. 
minutissima Prevot, Ann. Sci. Nat., 
Ser. Bot., 15, 1933, 134; Murray, in 
Manual, 5th ed., 1939, 288.) From 
Latin, very tiny. 

Differs from Veillonella gazogenes 
only in that the usual carbohydrates 
favor growth and that the gas formed is 
not absorbed by sodium hydroxide and 
is not inflammable. 

Non-pathogenic for rabbits, guinea pigs 
or white mice (Oliver and Wherry). 

Source : Two strains isolated from a 
mixed infection in aphthous ulcers of the 
gingival and buccal mucosa of a case of 
postpoliomyelitic paralysis. 

2c. Veillonella gazogenes var. syzygios 
Murray. (Syzygiococcus scarlatinae Herz- 
berg. Cent. f. Bakt., I Abt., Ref., 90, 
1928, 575; Micrococcus syzygios scarlati- 
nae Herzberg, Cent. f. Bakt., I Abt., 
Orig., HI, 1929, 373; Micrococcus syzygios 
Bergey et al.. Manual, 3rd ed., 1930, 92; 
Veillonella alcalescens var. syzygios Pre- 
vot, Ann. Sci. Nat., Ser. Bot., 15, 1933, 
134; Murray, in Manual, 5th ed., 1939, 
288.) From Latin, yoked. 

Differs from Veillonella gazogenes only 
l)y its abilitj' to grow under an atmos- 
pheric pressure of 4 cm mercury, with the 
formation of H2S in small amounts by 
some strains, and the production of 
nitrites from nitrates. 

Source: Found by Herzberg in 30 per 
cent of normal mouths and in 100 per cent 
of saliva from scarlet fever patients. 



FAMILY LACTOBACTERIACEAE 305 

FAMILY VII. LACTOBACTERIACEAE ORLA-JENSEN. 

(Orla-Jensen, Jour. Bact., 6, 1921, 271 ; Streptobacteriaceae Bergey, Breed and 
Murray, Preprint, Manual, 5th ed., 1938, 71.) 

Long or short rods, or cocci which divide like rods in one plane only, producing 
chains, but never tetrads or packets. Non-motile except for certain cultures of 
streptococci. Gram-positive. Pigment production is rare; a few species form a 
yellow, orange, red or rusty brown pigment. Surface growth on all media is poor or 
absent. Some species are strictly anaerobic. Carbohydrates are essential for good 
development; they are fermented to lactic acid, sometimes with volatile acids, alcohol 
and CO2 as by-products (except for the non-fermenting Diplococcus magnus). Gela- 
tin is very rarely liquefied. Nitrate is not reduced to nitrite. Found regularly 
in the mouth and intestinal tract of man and other animals, dairy products, fer- 
menting vegetable juices. A few are highly pathogenic. 

Key to the tribes of family Lactobacteriaceae. 

I. Cocci occurring singly, in pairs and in chains. 

Tribe I. Streptococceae, p. 305. 

II. Rods occurring singly, in pairs and in chains. Individual cells may be very 

long or even filamentous. 

Tribe II. Lactobacilleae, p. 349. 

TRIBE I. STREPTOCOCCEAE TREVISAN. 

(I generi e le specie delle Batteriacee, 1889, 29.) 

Cells spherical or elongate, dividing in one plane only, usually occurring in pairs or 
chains. A few species are strict anaerobes; none grow abundantly on solid media. 
Carbohydrates and polyalcohols are changed either by homofermentation to lactic 
acid or by hetero fermentation to lactic and acetic acids, alcohol and carbon dioxide. 
Some pathogenic species grow poorlj^ without blood serum or other enrichment fluids. 
Catalase negative. 

Key to the genera of tribe Streptococceae. 

I. Parasites, growing poorly on artificial media. Cells usually in pairs, often 
elongated. Anaerobic species rarely in tetrads or small clumps. 

Genus I. Diplococcus, p. 305. 

II. Parasites and saprophytes. Normally forming short or long chains. Fer- 

ment glucose to lactic acid with practically no other acids or CO2. 

Genus II. Streptococcus, p. 312. 

III. Saprophytes. Form chains of cocci to short rods in plant juices and milk. 

Ferment glucose with the production of CO2, lactic acid, acetic acid and 
ethyl alcohol. Mannitol is formed from fructose. 

Genus III. Leuconostoc, p. .346. 

Genus I. Diplococcus Weichselbaum* 

(Weichselbaum, Wiener med. Jahrb., 82, 1886,483; Hyalococcus Schroeter, in Cohn, 
Kryptogamen Flora v. Schlesien, 1886, 152 ;Pseudodiplococcus Bonome, Cent. f. Bakt., 

* Revised by Prof. E. G. D. Murray, McGill University, Montreal, Canada, Sep- 
tember, 1938; anaerobic section adapted from papers by Dr. A. R. Prevot, Institut 
Pasteur, Paris, France, 1938; further revision b}^ Lt. Col. Elliott S. Robinson, M.C., 
Washington, D. C, January, 1944. 



306 MANUAL OF DETERMINATIVE BACTERIOLOGY 

2, 1888, 321; ? Pneumococcus Schmidlechner, Ztschr. f. Geburtshilfe u. Gynakol., 66, 
1905, 291; not Pneumococcus Arloing, Compt. rend. Acad. Sci., Paris, 109, 1889, 430; 
Mogallia Enderlein, Sitzb. Gesell. Naturf. Freunde, Berlin, 1917, 309.) From Greek 
diploos, double; kokkos, a grain or berry; M. L., a sphere. 

Cells usually in pairs, sometimes in chains or more rarely in tetrads or small clumps. 
Young cells Gram-positive. Parasites sometimes growing poorly or not at all on 
artificial media. Fermentative powers usually high, most strains forming acid from 
glucose, lactose, sucrose and inulin. The aerobic species are bile soluble while the 
anaerobic species are not bile soluble. 

The relationships of the strictly anaerobic diplococci placed in this genus by Prevot 
(Ann. Sci. Nat., S6r. Bot., 15, 1933, 140) to pneumococci are not yet entirely clear. 
The anaerobic species are included here in the hope that this arrangement will stimu- 
late research. 

The type species is Diplococcus pneumoniae Weichselbaum. 

Key to the species of genus Diplococcus. 

I. Aerobic, facultative. Bile soluble. 

1. Diplococcus pneumoniae. 
II. Strictly anaerobic. Not bile soluble. 

A. Greater than 1 micron in diameter. 
1. Carbohydrates not attacked. 

2. Diplococcus magnus. 

B. Not greater than 1 micron in diameter. 

1. Acid from glucose and lactose. 

a. Capsulated. Pathogenic. 

3. Diplococcus paleopneumoniae . 
aa. Not capsulated. Non-pathogenic. 

4. Diplococcus plagarum-helli . 

2. Acid from glucose, not from lactose. 

a. Grows on ordinary culture media. Non-pathogenic. 

5. Diplococcus constellatus . 
aa. No growth on ordinary culture media. Pathogenic. 

6. Diplococcus morbillorum. 

1. Diplococcus pneumoniae Weichsel- Trevisan, I generi e le specie delle 

baum. (Microbe septicemique du salive, Batteriacee, 1889, 34); Weichselbaum, 

Pasteur, Chamberland and Roux, Compt. Wiener med. Jahrb., 82, 1886, 485; Pneu- 

rend. Acad. Sci., Paris, 93, 1881, 159; moniemikrococcus or Pneumococcus, 

Micrococcus of rabbit septicemia. Stern- Frankel, Ztschr. f. klin. Medizin, 10, 

berg. National Board of Health Bull., 1886, 402; Bacillus septicus sputigenus 

Washington,:?, 1881, 781; Coccus lanceole, Fliigge, Die Mikroorganismen, 2 Aufi., 

Talamon, Communication a la Societe 1886, 262; Bacillus salivarius septicus 

anatom. de Paris, 58, 1883, 475; Micro- Biondi, Ztschr. f.Hyg.,£, 1887,195;DzpZo- 

coccus pyogenes tenuis Rosenbach, coccus lanceolatus sive capsulatus Foa and 

Mikroorganismen bei den Wundinfek- Bordoni-Uffreduzzi, Archivio per le Sci. 

tionskrankheiten des Menschen, 1884, 30 Med., 11, 1887, 387; Streptococcus 

(see Neumann, Cent. f. Bakt., 7, 1890, lanceolatus pasteuri Gamaleia, Ann. Inst. 

177); Micrococcus pasteuri Sternberg, Fsist., 2, 1SS8, ■i42; Streptococcus layiceo- 

Trans. Pathol. Soc. of Philadelphia, 12, latus Gamaleia, ibid., 443; Klebsiella 

1885, 162 (not Micrococcus pasteuri salivaris Trevisan, I generi e le specie 



FAMILY LACTOBACTERIACEAE 



307 



delle Batteriacee, 1889, 26; Micrococcus 
rosenhachii Trevisan, ibid., 33; Micro- 
coccus pTjogenes-ienuis De Toni and 
Trevisan, in Saccardo, Sylloge Fungorum, 
8, 1889, 1031 ; Micrococcus pneumoniae 
crouposae Sternberg, Cent. f. Bakt., 12, 
1892, 53; Diplococcus lanceolatus capsu- 
latus Kruse and Pansini, Ztschr. f. Hyg., 
11, 1892, 335; Diplococcus lanceolatus 
incorrectly ascribed to Frankel by 
Binaghi, Cent. f. Bakt., I Abt., 22, 1897, 
278; Micrococcus tenuis Migula, Sj^st. d. 
Bakt., 2, 1900, 193; Bacterium pneu- 
moniae Migula, ibid., 347; Bacterium 
salivarium Migula, ibid., 379; Strepto- 
coccus pneumoniae Chester, Man. 
Determ. Bact., 1901, 63; Micrococcus 
lanceolatus Longcope, Jour. Med. Res., 
7 (N.S. 2), 1902, 220; Pneumococcus 
lanceolatus Schmidlechner, Ztschr. f. 
Geburtshilfe u. Gynakologie, 56, 1905, 
291; Pneumococcus pneumoniae Fried, 
Jour. Exp. Med., 57, 1933, 111.) From 
Greek pneumonia, inflammation of the 
lungs. 

Monas pulmonale Klebs (Arch. f. 
exper. Path. u. Pharmakol., 4, 1875, 472) 
is inadequately described by Klebs and 
ought not to be regarded as identical 
with Weichselbaum's organism. 

Common name: Pneumococcus. 

The organisms occur as oval or spherical 
forms typically in pairs, occasionally 
singly or in short chains, 0.5 to 1.25 
microns. The distal ends of each pair 
of organisms tend to be pointed or lancet- 
shaped. Encapsulated. Non-motile. 
Young cells, Gram-positive. 

Gelatin stab: Filiform or beaded 
growth. No liquefaction. 

Infusion agar colonies: Small, trans- 
parent, grayish, with entire margin. 
Elevation high convex, glistening, mucoid 
to watery. 

On blood agar, the colonies are elevated 
at the center with concentric elevations 
and depressions. Hemolysis usually 
slight but often marked in anaerobic 



culture; methemaglobin formation with 
green zone around colony. 

Beef heart infusion broth : Uniform 
turbidity with variable amount of 
sediment. 

Addition of glucose, serum, whole 
blood or ascitic fluid enhances growth. 

Meat extract media : Growth irregular, 
usually poor if any. 

Inulin serum water : Usually acid with 
coagulation. 

Litmus milk: Usually acid with 
coagulation . 

Potato: No growth. 

Whole bile or 10 per cent solutions of 
sodium taurocholate or sodium glyco- 
cholate added to actively growing broth 
cultures will dissolve the organisms. It 
is customary to use from 0.1 to 0.5 ml of 
bile for each 0.5 ml of culture. 

Aerobic, facultative. 

Optimum temperature 37°C. Usually 
no growth at 18° to 22°C. 

Optimum initial pH 7.8. 

Source : Sputum, blood and exudates in 
pneumonia ; cerebrospinal fluid in menin- 
gitis; mastoiditis; otitis media; peri- 
tonitis; empj^ema; pericarditis; endo- 
carditis; arthritis; saliva and secretions 
of respiratory tract in normal persons. 
Commonest cause of lobar pneumonia. 

Habitat : The respiratory tract of man 
and animals. 

At present, thirty-one types of Diplo- 
coccus pneumoniae are recognized on the 
basis of serological reactions, chiefly the 
Neufeld "Quellung" phenomenon as in- 
duced by type-specific immune rabbit 
serums. Following the description of 
Pneumococcus 1 by Neufeld and Handel 
(Arb. a. d. k. Gesundheitsamte, 34, 1910, 
293), Dochez and Gillespie (Jour. Amer. 
Med. Assoc, 61, 1913, 727) divided the 
species into Types 1, 2, 3 and a heterog- 
enous group 4; Cooper, Edwards and 
Rosenstein (Jour. Exp. Med., 49, 1929, 
461) separated Types 4 to 13 from the 
strains previously designated as group 4, 
and later Cooper, Rosenstein, Walter and 
Peizer (Jour. Exp. Med., 65, 1932, 531) 



308 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



continued the classification to Type 32. 
Due to marked cross-reactions, it was 
subsequently decided that Type 6 was 
identical with Type 26, and that Types 
15 and 30 were identical. This resulted 
in the deletion of the Cooper Types 26 
and 30, thus leaving thirty of the original 
thirty-two types. Type 33 (Wilder) has 
been described by Walter, Blount, Beat- 
tie and Cotler (Jour. Inf. Dis., 66, 1940, 
181) as a distinct type; sufficient recogni- 
tion has been accorded to justify the 
acceptance of this type, thereby making 
a total of thirty-one types of the species. 
In a still more recent publication, Wal- 
ter, Guevin, Beattie, Cotler and Bucca 
(Jour. Immunol., J!^l , 1941, 279) recom- 
mend the addition of nine new types and 
eight subtypes. These, together with 
new strains reported by Ivauff'mann, 
March and Schmith (Jour. Immunol., 
S9, 1940, 397), if eventually recognized, 
would make a total of fifty-five types. 
Eddy still more recently, taking into 
account all known types, raises the num- 
ber of recognized types to seventy five 
(U. S. Public Health Repts., 59, 1944, 
449-468) . 

Note 1. Streptococcus mucosus How- 
ard and Perkins. (Howard and Perkins, 
Jour. Med. Res., 6 (N.S. I), 1901, 174; 
Diplococcus capsulatus incorrectly at- 
tributed to Frankel by Binaghi, Cent. f. 
Bakt., I Abt., 22, 1897, 273; Strepfococcus 
tmicosus Schottmiillor, Miinch. med. 
Wchnschr., 50, 1903, 909; Streptococcus 
lanceolatus var. mucosus Park and Wil- 
liams, Diplococcus lanceolatus var. 7nu- 
cosus Park and Williams, Diplococcus 
mucosus Park and Williams, Pneumo- 
coccus mucosus Park and Williams, 
Jour. Exp. Med., 7, 1905, 411; Strepto- 
coccus mucosus capsulatus Buerger, Cent, 
f. Bakt., I Abt., ^U 1906. 314.) This 
organism is no longer recognized as a 
separate species. Dochez and Gillespie 
(Jour. Amer. Med. Assoc, 61, 1913, 727), 



Wirth (Cent. f. Bakt., I Abt., Orig., 102, 
1928, 40) and others have established the 
identity of strains of this group as Diplo- 
coccus pneumoniae, Type 3. 

Buerger (Cent. f. Bakt., I Abt., Orig., 
41, 1906, 314) lists the following capsu- 
lated closely related streptococci : Strep- 
tococcus involutus Kurth, Arb. a. d. k. 
Gesundheitsamte, 8, 1893, 449 {Diplo- 
coccus involutus Winslow and Winslow, 
The Systematic Relationships of the 
Coccaceae, New York, 1908, 131) ; Strep- 
tococcus aggregatus Seitz, Cent. f. Bakt., 
I Abt., 20, 1896, 854; Streptococcus cap- 
sulatus Binaghi, Cent. f. Bakt., I Abt., 
22, 1S97, 273; Streptocoque aureole, Le 
Roy des Barres and Weinberg, Arch, de 
Med. exper. et d'anat. pathol., 2, 1899, 
399; Leuconostoc hominis Hlava, Cent. f. 
Bakt., I Abt., Orig., 32, 1902, 263. 

Note 2. Pneumococci, regardless of 
serological type, manifest three chief 
culture phases (or stages): Mucoid, 
Smooth, and Rough. The Mucoid (M) 
form corresponds to that previously desig- 
nated as Smooth (S) and represents the 
typical phase of the species; Smooth (S) 
supercedes the earlier term Rough (R); 
and the present Rough (R) form is a rela- 
tively newly-described variant. The 
most frequently observed dissociative 
trend is M -^ S ^ R. Serological types 
are recognizable only in the Mucoid form 
due to the presence of type-specific poly- 
saccharides in the capsular material ; 
both Smooth and Rough forms are devoid 
of capsular material, but possess species- 
specific antigens common to all members 
of the species. Smooth and Rough forms 
are non-pathogenic, possess distinctive 
growth characteristics, and require spe- 
cial technic for accurate observations. 
The cultural characteristics given are 
those of the mucoid and smooth phases 
only, e. g., see growth in broth. 

'• t Diplococcus magnus Prevot. {Di- 
plococcus magnus anaerobius Tissier and 



* Anaerobic section reviewed by Dr. Ivan C. Hall, New York, N. Y. 

t These anaerobic diplococci and streptococci, many of which are putrefactive 



FAMILY LACTOBACTEKIACEAE 



309 



Martelly, Ann. Inst. Past., 16, 1902, 885; 
Pr^vot, Ann. Sci. Nat., S^r. Bot., 15, 
1933, 140.) From Latin magnus, large. 

Large spheres: 1.5 to 1.8 microns, 
usually in pairs, sometimes occurring 
singly, in small clumps or very short 
chains. Gram-positive. 

Gelatin: Growth slow, scanty. No 
liquefaction. 

Deep agar colonies : After 24 hours at 
37°C, lenticular, whitish, granular; mar- 
gin finely cut. No gas produced. 

Broth: Turbid, clearing in 4 or 5 days 
resulting in a viscous mass similar to the 
zoogloea which Clostridium hifcrmenians 
forms. 

Peptone water: Slight turbidity. 
Indole not formed. 

Milk: Unchanged. 

Fibrin not digested. 

Sterilized urine : Turbid in 3 tu 4 days. 
The urea is attacked forming (NH4)2C03. 

Proteoses : Digested and disintegrated 
forming (NH4)2C03 with the liberation 
of NH3. 

Carbohydrates not attacked. 

Optimum pH 7.0. Limits of pH 5.5 
to 8.5. 

Temperature relations : Optimum 37°C. 
Grows from 18° to 37°C'. Killed in five 
minutes on boiling or in half an hour 
at 60°C. 

Non-pathogenic. 

Strict anaerobe. 

Distinctive characters: Large size; 
very marked alkalinizing power. 

Source : Isolated by Tissier and 
Martelly {loc. cil.) from putrefying 
butcher's meat. Isolated by Prevot 
{loc. cit.) from a case of acute ap- 
pendicitis. 



Habitat : Human digestive tract. 
Very common on butcher's meat in the 
process of putrefaction. Probably occurs 
in household dust. 

3. Diplococcus paleopneumoniae 

Prevot. (An anaerobic pseudopneumo- 
coccus, Rist, These de Paris, 1898; Der 
Frankelsche Diplococcus, Bolognesi, 
Cent, f . Bakt., I Abt., Orig., 43, 1907, 113 ; 
Prevot, Ann. Sci. Nat., Ser. Bot., 15, 
1933, 143.) From Greek paleus, old and 
pneumonia, inflammation of the lungs. 

Spheres: About 0.7 to 1.0 micron, oc- 
curring in pairs, rarely occurring singly 
or in very short chains. Capsulated. 
Gram-positive. 

Gelatin: No liquefaction. 

Deep agar colonies : Probably len- 
ticular. 

Agar slant colonies : Round, raised, 
transparent, dew-drop. 

Broth : Opalescent turbidity which 
settles as a rather abundant, powdery, 
flocculent precipitate. No gas produced. 

Glucose or lactose broth: Rapid, 
abundant growth. 

Peptone water (2 per cent) : Very slow 
development. After 4 or 5 days at 37°C 
growth verj' poor. 

Milk: Good growth. Partial coagu- 
lation. 

Blood agar: Very rapid, abundant 
growth. 

Acid from glucose and lactose. 

Temperature relations : Optimum 37°C. 
No growth at 20°C nor at 42°C. Killed 
at 55°C. 

Pathogenic. 

Strict anaerobe. 

Distinctive characters : Resembles 



and gas-forming, seem to us so different from the fermentative microaerophilic 
diplococci, streptococci, leuconostocs and lactobacilli that we beleive they should 
be placed in genera and in a family separate from Lactobacteriaceae . Prevot in a 
discussion (Ann. Inst. Past., 67, 1941, 87) that has just reached us (Oct., 1945) 
recognizes this difference in physiology. He would solve the difficulty by return- 
ing the fermentative diplococci and streptococci to the family Coccaceae because 
of resemblances in morphology which do not seem to us to be fundamental — The 
editors. 



310 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



Diplococcus pneumoniae but is a strict 
anaerobe ; highly pathogenic . 

Source: Isolated by Rist (loc. cit.) 
from an osseous abscess; by Bolognesi 
(Joe. cit.) from lesions of pleuropneu- 
monia. 

Habitat : Buccal-pharyngeal cavity of 
man and rodents. 

4. Diplococcus plagartim-belli Prevot. 
(Diplococcus from septic wounds, Adam- 
son, Jour. Path, and Bact., 22, 1919, 393; 
Prevot, Ann. Sci. Nat., Ser. Bot., 15, 
1933, 157.) From Latin plaga, wound; 
belluni, war. 

Spheres: 0.6 to 1.0 micron, occurring 
in pairs of unequal size or in short chains. 
Gram-positive. 

Gelatin: No liquefaction. 

Deep agar colonies : Appear after 24 to 
48 hours, gradually increasing in size to 
2 mm in diameter; lenticular, regular, 
almost transparent. Gas not produced, 
even in glucose agar. 

Broth: Growth precipitates in 5 or 6 
days. No gas produced. 

Indole not formed. 

Milk : Strongly acidified and coagu- 
lated in 2 to 3 days. 

Serum not digested. 

Acid but not gas from glucose, mal- 
tose, lactose and sucrose. Xo acid from 
mannitol. 

Temperature relations : Optimum 
37°C. Not always killed in half an hour 
at 80°C. 

Non-pathogenic . 

Strict anaerobe. 

Source : Si.xteen strains isolated from 
fifty -one cases of septic war wounds. 

Habitat: Common in septic wounds. 

5. Diplococcus constellatus Prevot. 
(Compt. rend. Soc. Biol. Paris, 91, 1924, 
426.) From Latin constellatus, studded 
with stars. 

Description in part from Prevot, Ann. 
Sci. Nat., Ser. Bot., 15, 1933, 158. 

Spheres : 0.5 to 0.6 micron, occurring in 
pairs and tetrads, rarely in very short 



chains, never in clusters. Gram- 
positive. 

Gelatin: Good growth. No lique- 
faction. 

Deep agar colonies : At first very small, 
lenticular, biconvex, thick, opaque, 
yellowish, 0.5 to 1.5 mm in diameter. 
Each colony surrounded by many small 
satellite colonies visible microscopically. 

Broth: Growth slow, poor. After 48 
hours a slight homogenous turbidity 
which quickly clears, leaving a slight 
powdery sediment. Neither gas nor odor 
produced. 

Glucose broth : Growth rapid, 
abundant. 

Proteins not attacked. 

Blood broth: Good growth. No 
hemolysis. 

Milk: Poor growth. No change. 

Peptone water: Good growth. Not 
acidified. Indole not formed. 

Neutral red broth unchanged. 

Acid but not gas from glucose, 
arabinose. Slightly acid from glycerol. 
No acid from lactose, inulin, mannitol or 
dulcitol. 

Optimum pH 6.0 to 8.0. 

Optimum temperature 37°C. Feeble 
growth at 22°C. Not thermo-resistant. 

Strict anaerobe. 

Distinctive character : The micro- 
scopic appearance of agar colonies each of 
which is surrounded by a constellation of 
satellites. 

Source : Isolated from a case of chronic, 
cryptic tonsillitis. Later isolated from 
pus in acute appendicitis. 

Habitat: Digestive tract, especially 
the lymphoid tissues, as tonsils and 
appendix. 

6. Diplococcus morbillorum Prevot. 

(Diplococci from cases of measles, 
Tunnicliff, Jour. Amer. Med. Assoc, 68, 
1917, 1028; Diplococcus rubeolae Tunni- 
cliff, Jour. Inf. Dis., 52, 1933, 39; Prevot, 
Ann. Sci. Nat., Ser. Bot., 15, 1933, 148; 
original name withdrawn by Tunnicliff, 



FAMILY LACTOBACTERIACEAE 



311 



Jour. Inf. Dis., 58, 1936, 1.) From Latin 
morbus, disease; M. L. vwrhilli, measles. 

Spheres : 0.6 to 0.8 micron, occurring in 
short chains, rarely in small masses. 
Gram-positive. 

This organism does not develop on 
ordinary culture media. The addition of 
fresh serum or ascitic fluid is necessary. 

Gelatin : No liquefaction. 

Serum agar colonies : Very small, punc- 
tiform, appearing after 5 to 22 days. No 
gas produced. 

Glucose agar containing ascitic fluid 
and blood : Colonies are slightly larger 
and appear more rapidly ; greenish. 

Blood agar colonies : Surrounded by a 
greenish halo. May be large and moist. 
Gas not produced. 

Broth: Very poor growth. 

Hemolysed blood broth: Growth fioc- 
culent, leaving the liquid clear. 



Milk: Unchanged by most strains. 
.Acidified and coagulated by four strains. 

Indole not formed. 

Bile : Not soluble in bile. 

Acid from glucose, sucrose and 
maltose. 

Temperature relations : Optimum 
37°C. Killed in 45 minutes at 57°C. 
Withstands — 2°C for two weeks. 

Strict anaerobe. Most strains become 
microaerophilic with transfers. 

Distinctive characters: Greenish col- 
onies on blood media; poor growth on 
ordinary media. 

Source : Isolated from the throat and 
blood in measles. 

Habitat: Nose, throat, eyes, ears, mu- 
cous secretions and blood from cases of 
measles. 



312 MANUAL OF DETERMINATIVE BACTERIOLOGY 

Genus II. Streptococcus Roseiibach.* 

(Rosenbach, Mikroorganismen bei Wundinfektionskrankheiten des Menschen, 1884, 
22; Arthrostreptokokkus Hueppe, Wiesbaden, 1886, 144; Sphaerococcus Marpmann, 
Erganzungshefte z. Cent. f. allg. Gesundheitspflege, S, 1889, 121 ; Perroncitoa, Babesia, 
Schuetzia Trevisan, I generi e le specie delle Batteriacee, 1889, 29; Lactococcus Bei- 
jerinck, Arch, neerl. d. sci. exactes, Ser. 2, 7, 1901, 213; Hypnococcus Bettencourt et 
al.. Cent. f. Bakt., I Abt., Orig., 35, 1904, 55; Myxococcus Gonnermann, Oester. u. 
Ungar. Ztschr. f. Zuckerind. u. Landwirtsch., 36, 1907, 883; not Myxococcus Thaxter, 
Bot. Gaz., 17, 1892, 404; Melococciis Amiradzibi, Med. Zurn., 4, 1907, 309; Diplo- 
slreptococcus v. Lingelsheim, in Kolle and Wassermann, Handb. d. path. Mikroorg., 
2 Aufl., 4, 1912, 494; ? Brachybacterium Troili-Petersson, Cent. f. Bakt., II Abt., 11, 
1903, 138; Pseudostrephis Enderlein, Sitzb. Gesell. Naturf. Freunde, Berlin, 1917, 309; 
Planostreptococcus Meyer, Die Zelle der Bakterien, Jena, 1912, 4; Streptus Enderlein, 
Sitzber. Gesell. Naturf. Freunde, Berlin, 1930, 104; Peptostreptococcus Kluyver and 
Van Niel, Cent. f. Bakt., II Abt., 94, 1936, 391.) From Greek streptus, flexible or 
pliant; Greek kokkos, a grain or berry; M. L., a sphere. 

Cells spherical or ovoid, rarely elongated into rods, occurring in pairs, or short or 
long chains, never in packets or zoogloeal masses. Capsules are not regularly formed, 
but become conspicuous with some species under certain conditions. Gram-positive, 
some species decolorizing readily. A few cultures produce a rusty red growth in deep 
agar stab, or a yellow or orange pigment in starch broth. Growth on artificial media is 
slight. Agar colonies are small. Surface colonies are translucent. Colonies may 
be effuse, convex or mucoid. Some species are aided by the addition of native proteins. 
Mostly facultative anaerobes, with little surface growth in stab cultures. A few are 
strict anaerobes. Some of the latter attack proteins with production of gas and foul 
odors. Carbohydrate fermentation by all others is homofermentative, with dextro- 
rotatory lactic acidas the dominant product, while volatile acids, other volatile products 
and CO2 are either absent or produced in very small amounts. Inulin is rarely at- 
tacked. Nitrate is not reduced to nitrite. Not soluble in bile. Common wherever 
organic matter containing sugars is accumulated. Regularly in the mouth and 
intestine of man and other animals, dairj- products, fermenting plant juices. Some 
species are highly pathogenic. 

The type species is Streptococcus pyogenes Rosenbach. 

Note : The classification of streptococci is beset with many difficulties and it seems 
advisable for the present to accept only such described species about which there is 
reasonable agreement. With present knowledge, many species which have been sepa- 
rated can justifiably be considered as identical with older species and have been 
treated as such here. The descriptions of certain other species do not permit their 
exact identification now and they have been classed as invalid names with no present 
significance. It is admitted there are grounds for belief that more than one species 
may be included in certain of the species described here, but the onus of proof lies 
with the investigators interested in them. It is hoped that the simplification intro- 
duced will prove useful as a starting point for the more exact differentiation and 
description of the species of Streptococcus. The general arrangement used is in 

* Revised by Prof. E. G. D. Murray, McGill University, Montreal, Canada, in con- 
sultation with Prof. G. J. Hucker, New York State Experiment Station, Geneva, New 
York and Prof. J. M. Sherman, Cornell University, Ithaca, New York, June, 1938; 
further revision by Prof. J. M. Sherman, February, 1944. 



FAMlL,Y LAClOBACTil^KlACEAJlJ 313 

harmony with the suggestions made b}^ Hucker (Proc. 2nd Internat. Cong, for Micro- 
biology, London, 1936, 127) and Sherman (Bact. Reviews, 1, 1937, 3). 

Serological reactions are included as far as possible in the descriptions but the true 
significance of these methods is not known and on that account they are not stressed in 
the primary classification. 

Throughout the history of this genus motile streptococci have been reported oc- 
casionally (e.g., Streptococcus herbarum Schieblich, Cent. f. Bakt., I Abt., Orig., 134, 
1932, 269; Koblmuller, Cent. f. Bakt., I Abt., Orig., 133, 1934, 310; Stolting, Uber die 
Streptokokken des normal reifenden Tilsiter Kases. Inaug. Diss., Kiel, 1935, 51; 
Pownall, Brit. Jour. Exp. Path., 16, 1935, 155) but it is not known whether these con- 
stitute definite species or whether (Leveison, Ann. Inst. Past., 60, 1938, 93) motile in- 
dividuals occasionally appear in ordinarily non-motile species. 

The anaerobic streptococci have not been sufficiently studied to be sure whether 
they should be included in the genus Streptococcus or given separate generic rank. 
Their metabolic processes seem reason for the latter view. The descriptions given 
are taken from Prevot (Ann. Sci. Nat., Ser. Bot., 15, 1933, 23). 

The material is arranged accordingly in three categories : A key and complete de- 
scriptions have been prepared for clearly defined species, species of uncertain taxo- 
nomic relationships have been placed in Appendix I with their necessarily incomplete 
descriptions, while even less valid and unidentifiable species are merely listed in 
Appendix II. 

Key to the species of genus Streptococcus. 

I. Facultative anaerobic species. 

A. Pyogenic group. No growth at 10°C. No growth at 45° C. Generally 

beta hemolytic. Generalh^ do not curdle litmus milk and reduce lit- 
mus slowly if at all. Mannitol and glycerol generally not fermented. 
Not tolerant of 0.1 per cent methylene blue, 6.5 per cent NaCl and pH 
9.6. Produce ammonia from peptone. 

1. Sodium hippurate not hydrolyzed. 

a. Lactose fermented. 

b. Sorbitol not fermented but trehalose fermented. Lancefield 
Group A. 

1. Streptococcus pyogenes. 

bb. Sorbitol fermented and trehalose not fermented. Lancefield 
Group C. 

2. Streptococcus zooepidemicus . 

aa. Lactose may or may not be fermented. Lancefield Group C. 
b. Trehalose not fermented. 

3. Streptococcus equi. 
bb. Trehalose fermented. 

4. Streptococcus equisimilis. 

2. Sodium hippurate hydrolyzed. Lancefield Group B. 

5. Streptococcus agalactiae. 

B. Viridans group. No growth at 10°C. Growth at 45°C (few exceptions in 

Streptococcus mitis). Reduce litmus after curdling litmus milk ; sorbitol 
and glycerol generally not fermented; mannitol rarely. Not tolerant of 
0.1 per cent methylene blue, 6.5 per cent NaCl or pH 9.6. Not beta 



314 MANUAL OF DETERMINATIVE BACTERIOLOGY 

hemolytic (though they may be under anaerobic conditions) but show 
varying degrees of greening of blood. Do not produce ammonia from 
peptone (few exceptions in Streptococcus mitis). 

1. Lactose is fermented. 

a. Do not grow at 50°C. Greening or indifferent in blood agar. Raf- 

finose, inulin, salicin and dextrin generally fermented. Esculin 

generally attacked. Growth with 2 per cent NaCl. 

b. Do not survive 60°C for 30 minutes. Starch not hydrolyzed. 

Not tolerant of bile. 

c. Mucoid colonies produced on sucrose and raffinose media. 

6. Streptococcus salivarius. 

cc. Colonies not mucoid on sucrose or raffinose media. Inulin 
not fermented. 

7. Streptococcus mitis. 

bb. Survives 60°C for 30 minutes. Starch is hydrolyzed except 
by variety inulinaceus. Tolerant of bile. 

8. Streptococcus bovis (and varieties). 

aa. Grows at 50°C. No action on blood. Esculin not attacked. Raf- 
finose, inulin, salicin and dextrin not fermented. No growth in 
2 per cent NaCl. 

9. Streptococcus thermophilus . 

2. Lactose not fermented. Tolerant of bile. 

10. Streptococcus equinus. 

C Lactic group. Growth at 10°C. No growth at 45°C. Reduce litmus prior 
to curdling of litmus milk. Sorbitol and glycerol not fermented. Not 
beta hemolytic. Tolerate 0.1 per cent methylene blue, but do not 
tolerate 6.5 per cent NaCl or pH 9.6. 

1. Maltose and dextrin fermented. Ammonia produced from peptone. 

Growth at 40°C. Group N of Shattock and Mattick. 

11. Streptococcus lactis. 

2. Maltose and usually dextrin not fermented. Ammonia not produced 

from peptone. No growth at 40°C. 

12. Streptococcus cremoris. 

D. Enterococcus group. Growth at 10°C. Growth at 45°C. Usually reduce 
litmus prior to curdling litmus milk. Sorbitol, glycerol and mannitol 
generally fermented. May or may not be beta hemolytic. Tolerate 
0.1 per cent methylene blue, 6.5 per cent NaCl and pH 9.6. Ammonia 
produced from peptone. Lancefield Group D. 

1. Not beta hemolytic. 

a. Gelatin not liquefied. 

13. Streptococcus faecalis. 
aa. Gelatin liquefied. 

14. Streptococcus liquejaciens. 

2. Beta hemolytic. 

a. Mannitol and sorbitol fermented. 

15. Streptococcus zymogenes. 
aa. Mannitol and sorbitol not fermented. 

16. Streptococcals durans. 
IL Anaerobic species. 

A. Strict anaerobes. 

1. Gas and fetid odor produced. 

a. No general turbidity in broth. 



FAMILY LACTOBACTERIACEAE 



315 



b. Acid from maltose. 

17. Streptococcus anaerobius. 
bb. No acid from maltose. 

18. Streptococcus foetidus. 
aa. Turbidity in broth. 

b. No gas in Veillon's semisolid agar. No gas in peptone water. 

19. Streptococcus putridus. 

bb. Abundant gas in semisolid agar. Gas in peptone water. 

20. Streptococcus lanceolatus. 
2. No gas and no fetid odor produced. 

a. Milk not coagulated. 

21. Streptococcus micros. 
aa. Milk coagulated. 

b. Viscous sediment in broth. Semisolid agar colonies blacken 
with age. 

22. Streptococcus parvulus. 

bb. No viscous sediment in broth. Semisolid agar colonies do not 
blacken with age. 

23. Streptococcus intermedius, 
B. Microaerophilic. 

1. Strictly anaerobic on isolation, later microaerophilic. 

24. Streptococcus evolutus. 



1. Streptococcus pyogenes Rosenbach. 
(Fehleisen, Ueber Erysipel, Deut. Zeit. 
f. Chir., 16, 1882, 391; Erysipelkokken, 
Fehleisen, Die Aetiologie des Erysipels, 
Berlin, 1883; Rosenbach, Mikroorganis- 
men bei Wundinfektionskrankheiten des 
Menschens, 1884, 22; Streptococcus ery- 
sipelatos (sic) Rosenbach, ibid., 22; 
Micrococcus erysipelatis Zopf, Die Spalt- 
pilze, 2 Aufl., 1884, 86; Streptococcus 
erysipelatis Zopf, Die Spaltpilze, 3 
Aufl., 1885, 51 ; Streptococcus erysipelato- 
sus Klebs, Die Allg. Path., Jena, 1887, 
318; Micrococcus scarlatinae and Strepto- 
coccus scarlatinae Klein, Report of the 
Medical Officer of the Local Government 
Board for 1885-1886, No. 8, 1887, 85; 
Streptococcus conglomeratus Kurth, Arb. 
d. k. Gesundheitsamte, 7, 1891, 389; 
Streptococcus longus von Lingelsheim, 
Ztschr. f. Hyg., 10, 1891, 331 and 12, 
1891, 3C8; Streptococcus puerperalis 
Arloing, Septicemie puerperale, Paris, 
1892 (Jordan, Brit. Med. Jour, 1912, 1); 
Staphylococcus erysipelatosHesse , Ztschr. 
f. Hyg., 34, 1900, 3i7 ; Streptococcals longus 
pathogenes seu erysipelatos Schottmiiller, ' 
Munch, med. Wchnschr., 50, 1903, 909; 
Streptococcus longus hemolyticus Sachs, 
Ztschr. f . Hyg., 65, 1909, 466 ; Streptococcus 



longissimus Thalmann, Cent. f. Bakt., I 
Abt., Orig., 56, 1910, 248; Streptococcus 
hemolyticus Roily, Cent. f. Bakt., I Abt., 
Orig., 61, 1911, 87; Streptococcus epi- 
demicus Davis, Jour. Am. Med. Assoc, 
58, 1912, 1852; Jour. Inf. Dis., 15, 1914, 
378; ibid., 19, 1916, 236; Streptococcus 
hemolysans Blake, Jour. Med. Res., 36 
(N.S. 31), 1917, 116; Streptococcus pyo- 
genes haemolyticus Weisenbach, Compt. 
rend. Soc. Biol. Paris, 81, 1918, 819; 
Streptus scarlatinae Enderlein, Sitzber. 
Gesell. Naturf. Freunde Berlin, 1930, 
104; Streptococcus pyogenes var. scarla- 
tinae Hauduroy et al.. Diet. d. Bact. 
Path., Paris, 1937, 520.) From Greek 
pyon, pus; -genes, producing. 

Spherical or ovoid cells : 0.6 to 1 micron 
in diameter in cultures ; usually spherical 
in blood and inflammatory exudates ; oc- 
curring in chains or pairs. Capsules are 
variable, sometimes well developed and 
can be induced. Gram-positive. 

Gelatin stab: Growth slight; minute 
opaque colonies, little surface growth. 
No liquefaction. 

Nutrient agar: Small colonies, trans- 
lucent, convex, entire, slightly granular; 
colonies are variable ; confluent growth a 
thin transparent film; tendency for 



316 



MANUAL OF DETERMINATIVE BACTERIOLOGY 



colonies to remain discrete. Growth in- 
creased bj' addition of blood or native 
proteins. Pairs or short chains in surface 
growth and longer chains in condensation 
fluid of slants. 

Broth: Flocculent sediment of tangled, 
chains, supernatant broth often clear 
except in very young cultures. No 
pellicle. 

Potato: Very slight or no visible 
growth. 

Litmus milk : Acid, seldom curdled, 
and litmus reduced slowly or not at all. 

Acid from glucose, maltose, lactose, 
sucrose, salicin and trehalose. No acid 
from inulin, raffinose, arabinose, glycerol, 
mannitol, sorbitol or dulcitol. 

No hydrolysis of sodium hippurate, 
starch or esculin. 

Ammonia is produced from peptone. 

Temperature relations : Optimum tem- 
perature around 37°C. No growth at 
10°C or 45°C. Does not survive 60°C 
for 30 minutes. 

Chemical tolerance : Tolerates 2 per 
cent NaCl but not 4 per cent and 6.5 per 
cent. Final pH in glucose broth 4.8 to 
6.0; no growth at pH 9.6. Methylene 
blue 0.01 per cent and 0.1 per cent not 
tolerated and not reduced. Inhibited by 
bile but not soluble. 

Action on blood : Superficial and deep 
colonies cause hemolysis in blood agar, 
usually with a wide zone surrounding the 
colony, which may have a well-defined 
margin circumscribed by a zone of con- 
centrated hemoglobin ; the margin of the 
zone is ill-defined with some strains. 
Conditions defined by Brown (Rocke- 
feller Inst. Med. Res., Monograph 9, 
1919, 14) known as beta hemolysis. 
Soluble antigenic hemolysin of more than 
one kind produced in fluid cultures; in- 
fluenced by constitution of medium and 
presence of serum; one is oxygen-sensi- 
tive and another is oxygen-stable. 
Special precautions necessary for its 
demonstration (F. Smith, Jour. Bact., 
S.^, 1937, 585,603). 

Toxin: An erythrogenic toxin is pro- 



duced ; commonly associated with scarlet 
fever. Relatively thermostable. 

Fibrinolysin: Dissolves human fibrin 
but not fibrin of rabbit or ox blood. 
Markedly thermostable. 

Serology : Constitutes Group A of 
Lancefield (C substance ; polysaccharide) 
(Jour. Exp. Med., 57, 1933, 571). Types 
within the species are distinguishable 
(M substance; protein) ; 23 identified by 
Griffith (Jour. Hyg., 34, 1934, 542). 
Antigen common to the group (P sub- 
stance; nucleo-protein) also present in 
other Gram-positive cocci. 

Facultative anaerobe. Occasionally in 
primary culture from lesions, pus, etc. 
grows only in anaerobic culture. 

Source : Human mouth, throat and 
respiratory tract ; inflammatory exudates, 
blood stream and lesions in human disease 
of very varied character. Occasionally in 
milk and udder of cows. Dust in sick 
rooms, hospital wards and other contami- 
nated sites. 

Habitat : In human infections of many 
varied types. Occasionally in udder in- 
fections of cattle and perhaps other 
animal sources. 

2. Streptococcus zooepidemicus Frost 
and Engelbrecht. (Animal pyogenes. 
Type A of Edwards, Jour. Bact., 27, 
1934, 527; Frost and Engelbrecht, A 
Revision of the Genus Streptococcus, 
privately published, 1936, 3 pp. and The 
Streptococci, 1940, 25; Streptococcus pyo- 
genes animalis Seelemann, Deutsche 
tierarzt. Wchnschr., 50, 1942, 8 and 48.) 
From M. L., derived to mean animal 
cpideynicus. 

Morphology and general cultural char- 
acters resemble Streptococcus pyogenes. 
Mucoid colonies are common. Capsules 
are constantly demonstrable and promi- 
nent. Gram-positive. 

Gelatin stab: No liquefaction. 

Litmus milk: May be curdled, litmus 
not reduced or slowly after curdling. 

Acid from glucose, lactose and 
sorbitol. Acid may be produced from 
maltose, sucrose and salicin. No acid 



FAMILY LACTOBACTERIACEAE 



317 



from arabinose, trehalose, raffinose, in- 
ulin, glycerol or mannitol. 

Does not hydrolyze sodium hippurate, 
but starch and esculin may be split. 

Ammonia is produced from peptone. 

Temperature relations : Xo growth at 
10°C or at 4o°C. Does not survive 60°C 
for 30 minutes. 

Chemical tolerance : Tolerates 2 per 
cent XaCl but not 4 per cent and 6.5 
per cent. Final pH in glucose broth 
4.5 to 5.2. Xo growth at pH 9.6. Meth- 
ylene blue 0.01 per cent and 0.1 per cent 
not tolerated and not reduced. 

Action on blood: Beta hemolysis. 

Serology: Group C of Lancefield (loc. 
cit.). Cross precipitation with Strepto- 
coccus equi. 

Facultative anaerobe. 

Source: Blood stream, inflammatory 
e.xudates and lesions of diseased animals. 
Xot known from man. 

Habitat : Disease process of domestic 
and laboratory animals. (Horse: endo- 
metritis, foetus. Hog: septicemia. 
Cow: septicemia, metritis, foetus. 
Fowls : slipped tendon. Guinea pig: 
IjTnphadenitis. Rabbit: septicemia. 
Fox : pneumonia.) 

3. Streptococcus equi Sand and Jensen. 
{Bacillus adenitis equi Baruchello, Soc. 
Veter. de Venetie, Undine, 1886; Giornale 
di anatomia fisiologica et patologia degli 
animali domestici, Pisa, Sept., 1887 ; Sand 
and Jensen, Deuts. Ztschr. f. Tiermed., 

13, 1888, 436, dated December 27, 1887, 
Veterinary Congress, Copenhagen, 1887; 
sometimes incorrectly credited to Schtitz, 
Arch. f. wissens. u. prakt. Tierheilkunde, 

14, 1888, 172; Streptococcus cappelletti 
Chester, Manual Determ. Bact., 1901, 57 ; 
Streptococcus coryzae contagiosae equorum 
Schiitz, in Eisenberg, Bakt. Diag., 3 
Aufl., 1891, 270; Streptococcus schiitz, 
Bongert, in Kolle and Wassermann, 
Handb. d. path. Mikroorg., 2 Aufl., 6, 
■^913,208.) From Latin equus, horse. 

Possible synonyms : Streptococcus peri- 
ionitidis equi Hamburger, Cent. f. Bakt., 
I Abt., 19, 1896, 882 {Streptococcus peri- 



tonUidis Migula, Syst. d. Bakt., 2, 1900, 
21 ) ; Streptococcus pyogenes equi Hutyra, 
in Lehmann and Xeumann, Bakt. Diag., 
7 Aufl., 1927,221. 

Xote: Rivolta (Dei parassiti vegetali 
come introduzione alio studio delle malat- 
tie parassitarie e delle alterazione dell' 
alimento degli animali domestici. Turin, 
1873, 161 ) described chains of cocci in ade- 
nitis scrophula equorum, morbus glandu- 
losus. 

Holth, reported by C. O. Jensen 
(Handb. d. Serumtherapie u. Serum diag- 
nostik in d. Veterinar-med. (Klimmer- 
Wolff-Eisner) , 2, 1911 , 223 j , and Adsersen 
(Cent. f. Bakt., I Abt., Orig., 76, 1915, 
111 ) studied the fermentation reactions of 
Sand and Jensen's organism. Review of 
early literature given by Brocq-Rousseau, 
Forgeot and Urbain (Le streptocoque 
gourmeu.x. Revue de Pathologic Com- 
paree et d'Hygiene Generale, Paris, 
1925). 

Ovoid or spherical cells : 0.6 to 1 micron 
in diameter, sometimes in pus the long 
axis of the cells are transverse to the 
chain, and sometimes in the axis of the 
chain resembling str